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Annals 



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of the 



Missouri Botanical 



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Garden 




Volume VII 

1920 



With Seven ^iates and' Eigityit wo Figures 



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A ; i!i0 391 



Published quarterly by the Board of Trustees of the 
Missouri Botanical Garden, St. Louis, Mo. 

Entered as second-class matter at the Post Office at Lancaster, Pennsylvania, 

under the act of March 3 r 1879. 



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A branch of the PaMc l&fa*y 

St. Lomtt ftfat 









Annals 



of the 






Missouri Botanical Garden 






A Quarterly Journal containing Scientific Contributions 
from the Missouri Botanical Garden and the Graduate Labora- 
tory of the Henry Shaw School of Botany of Washington Uni- 
versity in affiliation with the Missouri Botanical Garden. 



George T. Moore 



Editorial Committee 



Benjamin M. Duggar 






Information 

The Annals of the Missouri Botanical Garden appears four times dur- 
ing the calendar year: February, April, September, and November. Four 



;titute a volume. 

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TUf. S%o.l 



STAFF 
OF THE MISSOURI BOTANICAL GARDEN 



Director, 

GEORGE T. MOORE. 



Benjamin M. Duggar, 

Physiologist, in charge of 
Graduate Laboratory. 

Hermann von Schrenk, 

Pathologist. 
Edward A. Burt, 

Curator of the Herbarium. 



Greenman 



Curator of the Herbarium 



Joanne L. Karrer, 

Research Assistant. 

Katherine H. Leigh, 

Secretary to the Director 



Nell C. Horner, 

Editor of Publications. 



BOARD OF TRUSTEES 
OF THE MISSOURI BOTANICAL GARDEN 



President, 

EDWARDS WHITAKER. 

Vice-President, 

DAVID S. H. SMITH. 



Samuel C. Davis, 
Edward C. Eliot. 
George C. Hitchcock. 
Edward Mallinckrodt. 



Leonard Matthews. 
William H. H. Pettus 
Philip C. Scanlan. 
John F. Shepley. 



i 



EX-OFFICIO MEMBERS: 



W. Frank Carter, 

President of the Board of Public 
Schools of St. Louis. 

Frederic A. Hall, 

Chancellor of Washington University. 



Henry W. Kiel, 

Mayor of the City of St. Louis. 



George T. Moore, 

President of the Academy of Sci 
ence of St. Louis. 



Daniel S. Tuttle, 

Bishop of the Diocese of Missouri. 



Charles A. Roe, Secretary. 



TABLE OF CONTENTS 



PAGE 



Hydrogen Ion Concentration and the 
Composition of Nutrient Solutions in 
Relation to the Growth of Seed Plants 
• B. M. Duggar 1- 49 

Humidity in Relation to Moisture Imbi- 
bition by Wood and to Spore Germina- 
tion on Wood S. M. Zeller 51- 74 

On Carbohydrate Consumption by Azoto- 
bacter chroococcum E. R. Allen 75- 79 






d 



E. A. Burt 81-248 



The Thelephoraceae of North America. 
XII 

Studies in the Physiology of the Fungi. 
XI. Bacterial Inhibition by Metabolic 
Products W. H. Chambers 249-289 

The Nutritive Value of the Food Reserve 

in Cotyledons B. M. Duggar 291-298 

Titration Curves of Certain Liquid Cul- 
ture Media J. L. Karrer and R. W. Webb 299-305 

The Use of "Insoluble" Salts in Balanced 
Solutions for Seed Plants B. M. Duggar 307-327 



General Index to Volume VII 



329-333 






I 







Annals 



of the 



Missouri Botanical Garden 



Vol. 7 FEBRUARY, 1920 No. 1 



HYDROGEN ION CONCENTRATION AND THE COM- 
POSITION OF NUTRIENT SOLUTIONS IN RE- 
LATION TO THE GROWTH OF SEED PLANTS 



B. M. DUGGAR 



Graduate 



Washington University 

Introductory 



>/ 



By the use of the terms mineral nutrition or salt requirements 
of plants there is connoted a group of physiological processes 
and environmental conditions in which the mineral salts play 
an important r61e. Without entering into an elaborate discus- 
sion of these terms it may at least be pointed out that perhaps 
neither one is satisfactorily comprehensive. The last-men- 
tioned is vague, and the other inadequate, due to the fact that 
when plants are grown in a so-called nutrient solution, as is well 
known, it is not merely the nutrient r61e the effects of which are 
followed. The concentration of the solute molecules and com- 
ponent ions of the culture solution directly affect the turgor, or 
osmotic surplus; the proportions of the ions — particularly of the 
cations — influence the permeability relations, which, operative 
through the protoplasts, seem to be the fundamental consideration 
in certain "antagonism" phenomena; the composition of the salts 
employed determines the acid-alkali equilibrium, that is, the hy- 
drogen ion concentration, the influence of which is apparently 
most complex; and these and other possibilities affect ultimately 
growth, which, in part, of course, involves the incorporation into 
the living framework of the ions of the component salts. 



Ann. Mo. Bot. Gard., Vol. 7, 1920 



(i) 



542' >7 



[Vol. 7 
2 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



For several years 1 the writer has given attention, as occasion 
permitted, to the reaction of the medium as affecting growth in 
soil solutions, also its effect upon toxic action; but it is only 
within the past two years that opportunity has been taken from 
time to time to examine this factor critically and so far as prac- 
ticable in respect to physiologically balanced nutrient solutions. 

The illuminating contributions of Clark and Lubs ('17) on 
the colorimetric determination of the hydrogen ion concentra- 
tion of culture media for microorganisms, and of other biological 
fluids, effected so great an improvement in the simpler technique 
involving active acidity and alkalinity that there is now avail- 
able a convenient, rapid, and sufficiently accurate method of 
investigating the hydrogen ion relations in connection with the 
salt requirements of higher plants. On the other hand, the 
extensive contributions of Schreiner and Skinner ('10, '10a, '11, 
'12), Tottingham ('14), Shive ('15), McCall ('16), and many 
others, on salt requirements and the constitution of the mineral 
nutrient solution have made it possible to select, within certain 
limits, well-balanced solutions as points of departure. In the 
work here reported, therefore, the writer has not concerned him- 
self to any great extent with an investigation of the effects of 
variations in the proportions of the different salts involved in 
the nutrient solution. In this last-mentioned direction the 
recent literature represents a rapid advance both in technique 
and in result, and while, as will be pointed out later, the problem 
may not be finally solved, it has certainly been placed on a 

rational basis. 

The triangle-diagram scheme was first rendered of biological 
significance by Schreiner and Skinner, referred to above, and it 
was effectively employed by them in studies of the relations of 

plants to certain toxic agents which might exist in the soil and 
in studies of the ameliorating action of the nutrient ions, K, 
N0 3 , and P0 4 , upon such deleterious compounds. They like- 
wise investigated by this means the growth of wheat as affected 
by different ratios of phosphate, nitrate, and potassium. The 
same general diagram scheme has been perfected and advan- 

1 During the progress of these investigations the writer has had much assistance in 
the details of the culture work from various members of the graduate laboratory, to 
all of whom he its indebted, especially to Mrs. Emily Schroeder, Research Assistant, 
1918-19, and Mr. R. W. Webb, Research Assistant, 1919-20. 



1920] 



DUGGAE — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 3 

tageously applied by Tottingham in his elaborate account of 
nutrient solutions, and by Shive, McCall, and others in intensive 
analyses of physiological balance. The effect of variation in 
H-ion concentration may be approached finally in the same way. 
In this rather general survey of certain aspects of the subject, 
however, it has seemed that the factors are for the most part too 
complicated for most economical treatment by this method 
alone. 



Technique and Materials 

In the earlier work I selected for comparative studies on the 
influence of variation in H-ion concentration two solutions only : 
namely, a slight modification of Shive's R 5 C 2 , considered his 
best solution of optimal concentration, and a much modified 
Crone solution developed in this laboratory. Later there was 
added the R 6 d solution of Livingston and Tottingham, and 
ultimately some other combinations which seemed worthy of 
consideration. 

The partial volume-molecular proportions of the particular 
Shive solution employed are as follows: KH 2 P0 4 , 0.0180; Ca 
(N0 3 ) 2 , 0.0052; and MgS0 4 , 0.0150. Ferric phosphate in small 
amount is added, the partial concentration being 0.0044 gm. per 
liter of solution. This insoluble salt is used at such a low con- 
centration that its presence as a precipitate is scarcely percept- 
ible. I have employed the same salts in the same proportions, 
except that "soluble ferric phosphate" has been substituted for 
the insoluble iron salt in all cases not otherwise indicated. This 
substance is described as consisting of scales of ferric phosphate 
with sodium citrate, possibly as a single salt. The exact molec- 
ular composition of this iron-furnishing substance is unknown. 
From the description in the National Standard Dispensatory 
(1905) it will be seen that it is commonly made by the addition 
of 50 gms. ferric citrate and 55 gms. sodium phosphate, unef- 
floresced, to 100 cc. distilled water. It is barely possible that 
four salts are present, namely, sodium and ferric phosphate and 
sodium and ferric citrate. This salt combination possesses the 
advantage of solubility to a high degree, yielding what appears 
at first to be a true solution but seems in reality a colloidal solu- 
tion of high dispersity. Used in such extreme dilution as 



[Vol. 7 



4 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



indicated in the Shive solution, there is no noticeable precipita- 
tion, even on standing. There is every reason to believe that 
this substance is an excellent source of iron. The Shive solu- 
tion prepared in this way is designated solution A, and it is best 
to think of it as solution A, because, as developed later, Shive 
does not report the P H for his solutions, and this three-salt solu- 
tion may possess very different values in plant growth, depend- 
ing upon the grade or reaction of the monobasic phosphate 



employed 



Crone solution is made up as follows: water, 



1000 cc. ; KN0 3 , 1 gm. ; CaS0 4 , .5 gm. ; MgS0 4 , .5 gm. ; Ca 3 (P0 4 ) 2 , 
.25 gm.; and Fe 3 (P0 4 ) 2 .25 gm. In our solution we have halved 
the concentration of the first two salts, omitted the tricalcium 
phosphate, and substituted for the ferrous phosphate the " sol- 
uble iron phosphate " above mentioned. In fact, the solution as 
I have used it may no longer be called the Crone solution, but it 
was of interest in this work at the time, not because it was 
expected to compare favorably with solution A above, but rather 
because it differed widely in the combinations of the components 
and had afforded in my work very good, healthy growth. The 
partial volume-molecular proportions are as follows: KN0 3 , 
0.00495; CaS0 4 , 0.000726; MgS0 4 , 0.000526; and "soluble iron 
phosphate," 0.125 gm. per 1000 cc. It will be noted that this 
is in reality a four-salt solution differing also notably from the 
earlier solutions of Sachs, Knop, Pfeffer, and Meyer. Inasmuch 
as the iron salt supplies also the phosphate, this B solution 
contains a higher concentration of Fe than the usual culture 
solution. There is more or less precipitation of a light or slowly 
settling iron salt, when the combined solution is prepared. 
However, by adding the soluble ferric phosphate last and at the 
greatest dilution possible under the conditions a light precipitate 
only is formed, and this develops slowly in the form of suspen- 
sion films. It introduces no difficulties into the preparation of 
the solution, though perhaps renders its composition somewhat 
less definite. The films are so light that it is not difficult to 
remove uniform samples from the stock flasks. The osmotic 
value of this solution has not been determined, but it is obviously 
much less than that of the Shive solution, the latter being about 
1.75 atmospheres. 



1920] 

DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 5 



The solution of Livingston and Tottingham, here designated 
solution C, possesses the following partial volume-molecular 



concentrations: KN0 3 , 0.0216; Ca(H 2 P0 4 ) 2 , 0.0026; and 
MgS0 4 , 0.0150. Iron is supplied as in the case of solution A, 
the modified Shive solution. It is assumed that this solution 
has approximately the osmotic value of solution A above 

described. 

The methods of experimentation employed were in large 
measure those which I have described elsewhere ('11). "Well- 
seasoned" tumblers holding somewhat more than 250 cc. were 
used as culture vessels in all cases, the process of seasoning new 
tumblers consisting of filling them with a weak acid-dichromate 
cleaning solution and then steaming in the autoclave for one 
hour at 15 pounds pressure. Subsequently the tumblers were 
thoroughly washed and rinsed with distilled water. The dis- 
tilled water used here and in preparing the solutions was in some 
series from a Stokes still and in other series double distilled from 
glass. After transferring, with pipette or burette, the required 
amounts of each constituent stock solution to the tumbler, the 
volume was made up to 240 cc. In those cases in which Canada 
field peas were used the tumblers were covered with heavy paraf- 
fined paper made fast by rubber bands. Small holes were 
punched in the paper and the radicles of the seedlings inserted. 
Additional support for the growing seedlings was afforded by 
means of wire supports. In the case of both wheat and corn 
the seedlings were inserted into notches or holes in a paraffined 
cork, the latter just fitting the mouth of the tumbler. All 
cultures contained ten plants, and duplicate tumblers were 
arranged in every case not otherwise indicated. 

In most of the experiments here reported Merck's blue label 
chemicals have been employed, but under the conditions existing 
at the time it was not possible to be entirely uniform in this 
regard, and other standard reagents were used where necessary. 
No recrystallization or other purification method was applied 
to any salt employed in the culture solutions. Stock solutions 
of convenient concentration were prepared, and so far as prac- 
ticable every factor and procedure was made uniform, or com- 
parable, throughout. It soon became evident that the content 
of free phosphoric acid in the dihydrogen potassium phosphate 



[Vol. 7 
6 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



was not constant in the different makes and bottles of the 
reagent necessarily used, and a closer examination of this point 
as the work progressed indicated clearly that we were not secur- 
ing the uniformity of P H in the solution which we felt that we 
might reasonably expect. It was not anticipated that the 
standard as indicated by Sorensen ('09- 10) and others would be 
attained unless the reagent were guaranteed free from phos- 
phoric or other acids. The variation in P H is often too great 
to make it at all certain what is meant when this salt is desig- 
nated merely as acid potassium phosphate. One manufacturer 
who was appealed to on account of the variability referred to, 
assured the writer that it was not possible to furnish the salt 
free of phosphoric acid under the existing conditions. 

In this work the hydrogen ion concentration was invariably 
determined by the method of Clark and Lubs ('17), employing 
both their standard solutions and their indicators. The stand- 
ard solutions in this connection, however, were prepared with 
salts recrystallized two or three times, and the greatest care 
was given to every detail of the method. They were not con- 
trolled by the electrode method but by close comparison with 
the established effective ranges of the different indicators. esDe- 



indicators with 



When making 



comparisons there was arranged for each indicator a set of 
seasoned serological test-tubes each containing 5 cc. of any 
standard solution within the range of the particular indicator. 
The various sets were arranged on a rack with white paper 
background. From 3 to 5 drops of the indicator solutions were 
used. Inasmuch as many of the culture solutions employed 
were without color, sufficiently accurate determinations were 
readily made with the unaided eye. In the case of solution B, 
however, and likewise in certain cases referred to later, whe 
few algae appeared in the solutions, recourse was had to the 
method employing the colorimeter (Duggar and Dodge, '19) 
and later more especially to the micro-colorimeter (Duggar, '19). 
The seedlings used in various phases of the work were wheat 
(Triticum vulgare), corn (Zea Mays), and field peas (Pisum 
arvense). In all the work reported in tables i-vi, also xi- 
the wheat was of the variety Fultz, and that reported in tables 
vn-x was with a new variety, the Pacific Coast Blue Stem, 



a 




1920] _ 

DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS / 

supplied from the Plant Introduction Garden of the Bureau of 
Plant Industry at Chico, California. The latter had been found 
by Dr. H. S. Reed to be particularly good for solution culture 
work, and my experience is entirely confirmatory. The other 
seed were from selected but unnamed field varieties. 

The seed were immersed over night in running water, and 
then the peas and corn placed for germination on paraffined wire 
netting over pans of well-washed, moist sphagnum. They were 
covered with moistened paper toweling, over which was inverted 
other pans, though ample ventilation was provided. The wheat 
was treated in much the same way except that it was germinated 
over water frequently changed. As soon as the plumules 
emerged light was admitted. After properly placing the seed- 
lings, ten to each tumbler, they were left in diffuse light, in the 
room in which set up, for 12 to 24 hours in order to become 
better adjusted to the conditions before being installed in the 

greenhouse. 

In some of the earlier experiments a complete series was 

arranged under a single set of conditions, and in such cases the 
tumblers were placed upon a rotating table. In much of the 
later work, however, more than one set of conditions was in- 
volved, so that the use of rotating tables was not practicable, 
and under the circumstances special care was taken with prop- 
erly placing and spacing the tumblers on lattice tables, likewise 
shifting the order of the cultures so as to be wholly comparable. 

In all cases as the plants were harvested, the remains of the 
seed, or cotyledons, were cut away. This was found necessary 
inasmuch as otherwise a considerable error would be introduced 
in the weights of those cultures in which relatively little growth 
occurred; for it was in such plants that the seed were incom- 
pletely exhausted. On removal from solutions containing 
precipitates the roots were thoroughly washed, and in all cases 
quickly and uniformly dried of surface water on absorbent gauze. 

The criterion of growth on which stress is laid in this paper 
was total green weight. Other data included green weight of 
tops, average length of shoot (or leaf), dry weight, and general 
appearance — including root characteristics. Total green weight 
is, for the present purpose, entirely satisfactory and has the 
advantage in these relatively short-interval cultures of expressing 



c IVol. 7 

o ANNALS OF THE MISSOURI BOTANICAL GARDEN 



as truly as may be the growth and health of the plants. Dried 
leaves and withered tips count for little. Average length of 
leafy shoot is included in some of the tables, but this affords 
merely an index of stockiness or attenuation of the plants. 

The purpose in this work was to get results not only in respect 
to growth relations with variations in the H-ion concentration, 
but also, especially in the later work, to secure data in respect 
to the extent of change in the P H of the medium in which the 
plants had grown. The time interval which the plants were 
permitted to remain in the cultures between changes of solution 
was generally 6 to 7 days, but in certain cases referred to later 
this interval was diminished or increased for special reasons 
indicated in connection with the tabulated data. During any 
interval, however, distilled water was added as required. In 
general, it seemed to the writer that the practical value of a 
particular mineral nutrient solution should rest in part upon its 
capacity to furnish favorable growth conditions for a period not 
too limited, that is, the solution should possess among other 
favorable characteristics, if possible, the quality of resisting 
unfavorable change over a period of one week or even longer. 

Had the nutrient solutions used been merely solutions A and 
C, in which phosphate is supplied as the acid salts, it would 
have been simple to change the H-ion concentration towards 
neutrality by the use of various proportions of a dibasic salt. 
However, it was at the same time desired to increase the H-ion 
concentration, that is, to diminish the P H exponent in order to 
determine approximately the favorable P H limits. Moreover, 
changes in solution B could not be readily effected in the manner 
indicated without material changes in composition. It was 
therefore determined to use, in general, 0.1 n NaOH and approx- 
imately 0.1 molar H 3 P0 4 in shifting the H-ion concentration. 
Actually, the concentrations employed were 0.1 n NaOH and 
0.092 molar H 3 P0 4 . The stock solutions were so arranged as to 
concentration that the amounts of alkali or of acid introduced 
might first be diluted considerably with distilled water. In this 
way there was less danger of precipitating out the calcium salt, 
as insoluble calcium phosphate, for example, in solution A, 
before the critical P H was reached, also other possibilities of 
precipitation in the case of solution R. 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 9 



The problem was then to shift the hydrogen ion concentration 
of the solutions used without disturbing any more than necessary 
the composition of the medium. From preliminary tests of 
solutions A and B it was inferred that the chief interest might 
attach to the addition of alkali to solution A and of acid to 
solution B. The P H exponent of the former is low, but in the 
latter it more nearly approaches neutrality. It is seen that the 
active acidity of the Shive solution is due to dihydrogen potas- 
sium phosphate, and the relatively slight variation in the pro- 
portions of the other practically neutral salts may be assumed 
not to change materially the hydrogen ion concentration. In 

■ 

this sense, therefore, the 108 solutions tested by Shive are prob- 
ably nearly identical and determine, in respect to growth, the 
values of the proportions of the nutrient ions present in the 
solutions only in relation to this one value of P H . This value 
might, of course, be assumed to be most favorable under all 
environmental conditions, but it is equally possible that it is not. 
This is a factor which should be given most careful considera- 
tion in all mineral nutrient, or salt-balance, studies as well as in 
studies upon toxic action. 

As a convenient index to the content of the various cultures a 
simple scheme of notation has been devised, which, with very 
slight call upon the memory, enables one to see at a glance the 
constitution of the culture medium. This involves a system of 
letters and numerals as explained below. The initial letter of 
a culture refers to the general constitution of the nutrient solu- 
tion, and as stated above there are three such solutions. The 
modified Shive solution employed is designated A; the solution 
remotely based on the Crone formula is solution B; and the 
Livingston-Tottingham solution, C. The next letter is invari- 
ably a small letter and denotes the plant employed ; thus w desig- 
nates wheat; p, Canada field peas; and c, corn; while x is em- 
ployed where more than a single kind of plant is designated, or 
where no particular plant is specified. A culture index such as 
ApO, BpO, or CpO, etc., refers to the use of peas with the solu- 
tions mentioned without the addition of acid, alkali, or other 
constituent. When acid or alkali is added an inclined line repre- 
sents, in this case, neutrality, and when figures are given above 
the line mentioned they represent cc. of 0.092 molar phosphoric 



10 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



acid; and when given below they represent cc. of 0.1 n sodium 
hydroxide. Thus Ap/10 indicates the addition of 10 cc. of 

0.1 n NaOH to the A solution with peas. For the purpose of this 
paper we have considered molar phosphoric acid to be the sum of 
the values derived by titration against normal alkali, using as 
indicators both phenolphthalein and methyl orange. 

I have attempted to control or counterbalance, in a measure, 
the addition of phosphorus when added as phosphoric acid by 
the introduction in one or two cultures of an amount of phos- 
phorus in the form of the secondary salt equivalent to the acid 
added. At the same time, however, the addition of this salt 
necessarily shifts slightly the hydrogen ion concentration toward 
neutrality. The addition of sodium in the form of the alkali 
was in a measure controlled by adding as sodium sulphate an 
amount of this cation which is equal to that supplied in 10 cc. 
of alkali. As the work has progressed some other variations 
have been employed, notably the addition of solid calcium car- 
bonate (1 gm. per culture, 240 cc.) to solution B, likewise of solid 
aluminium hydroxide, and of kaolin, the addition of phosphate 
in part as the primary salt and in part as the secondary salt, 
also variations in the form and amount of phosphate employed, 
and significant changes in the proportions of solution B. Modi- 
neat ions of the culture indices denoting such changes will be 
explained as these are introduced. 



Experimental Data 

There are given in tables I, n, and in the results of the first 
tests conducted with wheat, corn, and peas respectively, using 
solutions A and B. In consulting these tables it is to be remem- 
bered that the " culture indices" are intended to afford briefly all 
necessary facts concerning the constitution of the solution, and 

an explanation of these has been made on p. 9, so far as the 
unmodified solutions and those containing additions of acid and 
alkali are concerned. In the tables referred to above, K/10 and 
K/20 represent respectively additions of K 2 HP0 4 to balance the 
amounts of phosphoric acid added in cultures with similar 
numerals; and Na indicates the addition of sodium sulphate to 
equal the quantity of gram atoms of Na in 10 cc. of n/10 NaOH. 
The wheat cultures were grown 17 days, the corn 21, and the 



1920] 



DTJGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



11 



peas 18. The solutions were renewed and water loss was sup- 
plied as previously noted. The determination of the hydrogen 
ion concentration "after" growth in the case of corn furnishes 
an index of the change in the solution after the plants had grown 
in it for 10 to 12 days, representing the final interval. In this 
particular case the intervals between changes of solution were 
made rather long, with the idea of emphasizing conditions. The 
experiments were begun early in November under favorable 
growing conditions. The daily evaporation rate from a stan- 
dardized spherical evaporimeter averaged 11 cc. per day. 



TABLE I 

(Series 1, Wheat) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH; 

SOLUTIONS A AND B 







Total 


Total 


Greatest 




No. 


Culture 
indices 


gr. wt. 
(gms.) 


dry wt. 
(gms.) 


length 
(cm.) 


Initial Ph 
of sol. 


1 


AwO 


4.92 


. 4077 


24.32 


4.0 


2 


Aw/5 


4.62 


0.3916 


23.89 


5.6 


3 


Aw/10 


5.20 


0.4135 


23.69 


5.9 


4 


Aw/15 


3.94 


. 3550 


21.24 


6.1 


5 


AwNa 


4.72 


. 3792 


25.93 


4.0 


6 


BwO 


4.52 


. 3545 


24.19 


7.0 


7 


Bw/5 


4.94 


. 3498 


23.36 


8.4 


8 


Bw/10 


5.83 


. 4090 


22.27 


9.0(+) 


9 


Bw5/ 


2.53 


. 2522 


16.48 


4.6 


10 


Bw 10/ 


0.69 


0.1176 


8.26 


3.2 


11 


Bw20/ 


0.67 


0.1134 


6.56 


3.0(-) 


12 


BwK 2 /10 


6.38 


. 4390 


26.34 


6.2 


13 


BwK 2 /20 


5.80 


0.3996 


25.94 


6.2 



From series i, table I, also fig. 1, it is seen that with wheat 
there is indication that the addition of a certain amount of 
alkali, shifting slightly the H-ion concentration, is beneficial, 
though Aw/5 is irregular. Wholly unexpected is the extent of 
the growth in certain of the B cultures. Introduced empirically, 
this solution has not only yielded well, but in their vigorous, 
green appearance these cultures rank highest. The addition of 
alkali to solution B is beneficial, possibly from the addition of 
the sodium ion or from the slight increase in the concentration 



12 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 




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9. + v. 6 

5 9 



3*4/ 3*/°/ ^Bx. 2 ^ Bx£, BxK so 

s, % 



JO 



3. -) 
11 



V 



e.z 6.0 



12 



13 



Fig. 1. Total green weight of wheat, com (5), and Canada field peas (5), in solu- 
tions A and B, with variations — chiefly in H-ion concentration. 



of the solution ; and even more favorable is the addition of small 
amounts of secondary phosphate. With corn, table 11, the 
shift towards alkalinity in the A solution affords slightly in- 
creased growth, but it is an important fact that there is a rapid 
falling off in the neighborhood of P H 6. This is approximately 
the H-ion concentration at which precipitation begins. With 
this crop, the B solution alone, solution B plus small amounts of 
alkali, and solution B with the addition of dibasic phosphate 
yield strikingly heavier than solution A. On the other hand, 
peas, under the conditions of these experiments, made maximum 
growth in the relatively acid solution A. 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



13 



TABLE II 

(Series 1, Corn) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH; 

SOLUTIONS A AND B 



No. 



1 

2 
3 
4 
5 
6 

7 
8 
9 
10 
11 
12 
13 



Culture 
indices 



AcO 

Ac/5 
Ac/10 

Ac/15 

AcNa 

BcO 

Bc/5 

Bc/10 

Be 5/ 

Be 10/ 

Be 20/ 

BcK 2 /10 

BcK 2 /20 



Total 


Gr. wt. 


Total 


gr. wt. 


of roots 


dry wt. 


(gms.) 


(gms.) 


(gms.) 



9.84 


2.91 


. 5452 


9.94 


2.74 


0.6139 


10.71 


2.43 


. 7642 


7.90 


2.25 




7.92 


2.56 


0.4652 


14.35 


4.22 


1.0198 


13.12 


2.70 


0.8102 


16.90 


3.95 


1.0271 


5.23 


1.17 


. 4606 


3.43 


0.70 


. 5387 


3.32 


0.41 


05617 


17.75 


3.94 


1.0170 


19.22 


4.51 


1 . 1881 



Greatest 
length 
(cm.) 



22.02 
22.64 
22.97 
20.36 
14.49 
25.79 
26.28 
29.24 
14.67 
10.54 
8.42 
31.84 
30.09 



P 



H 



Initial 



After gr. 



4.0 
5.6 
5.9 
6.1 
4.0 
7.0 
8.4 

9.0(+) 

4.6 

3.2 

3.0(-) 
6.2 

6.2 



5.4 
56 
59 
60 
5.2 
50 
8.0 
8.4 
4.0 
4.0 
4.0 
7.2 
6.6 



TABLE III 

(Series 1, Peas) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH; 

SOLUTIONS A AND B 



No. 



1 
2 
3 
4 
5 
6 
7 

8 
9 
10 
11 
12 
13 



Culture 

indices 



Total 
gr. wt. 

(gms.) 



ApO 
Ap/5 

Ap/10 

Ap/15 

ApNa 

BpO 

Bp/5 

Bp /10 

Bp5/ 

Bp 10/ 

Bp20/ 

BpK 2 /l0 

BpK 2 /20 



Gr. wt. 

of roots 

(gms.) 



Total 

dry wt. 
(gms.) 



Greatest 
length 

(cm.) 



19.47 


6.44 


17.37 


5.31 


17.71 


5.51 


17.10 


5.42 


14.77 


5.57 


15.72 


5.76 


15.86 


5.10 


14.10 


4.56 


8.27 


2.70 


5.85 


1.78 


3.47 


0.93 


17.69 


5.17 


15.39 


4.79 



1.1014 
0.9368 
1.0155 
1.0140 

. 8786 
. 9200 
0.8559 
. 7822 
. 6558 
0.5184 
0.3770 
. 9770 
. 8660 



27.84 
27.92 
29.42 
26.76 
25.60 
25.10 
27.40 
25.59 
15.83 
12.98 
7.59 
29.19 
27.18 



Initial P H 
of sol. 



4.0 
5.6 
5.9 
6.1 
4.0 
70 
8.4 

9.0(+) 

4.6 

3.2 

3.0(-) 

6.2 

6.2 



14 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 




Fig. 2. Total green weight of wheat, corn ($), and Canada field peas (j), in solu- 
tions A and B, with variations — chiefly in H-ion concentration. The base lines are 
drawn through cultures strongly acid, P H 3.4. 



A point of considerable interest, emphasized particularly by 
the results with peas, is that the range of most favorable growth 
with respect to hydrogen ion concentration differs materially 
with the constitution of the nutrient solution ; thus the addition 
of acid to the B solution, although shifting the H-ion concen- 
tration towards that of the A solution, exhibits a corresponding 
rapid diminution of the growth quantities. From these data 
alone it is not possible to formulate an explanation of the fact 
last mentioned, but it is probably related in part to ionic condi- 
tions, in part to the composition and state of aggregation of the 
iron and calcium particles, or to other, indetermined factors. 

It should be pointed out, that in examining the curve, fig. 1, 
and all subsequent curves, a horizontal, or base line, is drawn 
through the growth quantity representing generally the unmod- 
ified solution A, so that all cultures may be compared with this, 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



15 



TABLE IV 

(Series 2, Wheat) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 



No. 



1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 

16 

17 



Culture 
indices 



Total 


Total 


gr. wt. 


dry wt. 


(gms.) 


(gms.) 



AwO 

Aw/10 

Aw/20 

Aw/40 

AwNa 

AwM 

Aw 1/ 

Aw 3/ 

BwO 

Bw 1/ 

Bw2/ 

Bw5/ 

BwK 2 /10 

BwK /10 

BwCa 

Bw/5 

Bw/10 



4.39 


.545 


5.30 


.611 


4.56 


.533 


3.21 


.431 


4.86 


.556 


5.93 


.542 


5.21 


.569 


1.79 


.306 


6.07 


.503 


7.76 


.625 


7.82 


.657 


4.96 


.500 


7.60 


.623 


7.51 


.602 


8.67 


.810 


6.75 


.570 


7.08 


.606 



Greatest 
length 
(cm.) 



24.37 
24.75 
22.74 
19.80 
25.01 
23.04 
24.63 
16.38 
21.86 
23.96 
24.40 
21.81 
26.18 
2545 
25.32 
23.24 
24.49 



Initial P H 
of sol. 



3.4 
5.8 
6.3 



3.45 
5.4 

3.35 
3.15 
6.6 
6.4 

5.6 
3.2 
7.0 



7.3 
7.4 

8.6 



* Heavy precipitate. 



and the number of points in the curves above or below the base 
lines indicates for each crop, under the conditions reported, the 
relative increase or decrease in green weight. 

The change in the P H of the solutions occurring as a result of 
contact with the roots was followed only in the case of corn. 
As a rule, in the more acid solutions the reaction is shifted some- 
what towards neutrality, but irregularities occur in solution B, 
some of which may be related to changes not due to the inter- 
change of ions between roots and solution. 

The second series of experiments, the results of which are 
included in tables iv, v, and vi, also plotted in fig. 2, was carried 
through during late November and early December. The con- 
ditions were much the same in general as those prevailing during 
the earlier work. There was this difference, however, that while 
the first series was placed on latticed tables in greenhouses with 
proper spacing to provide for favorable and uniform condi- 



16 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 









TABLE 


V 










(Series 2, Corn) 






SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 


No. 


Culture 
indices 

• 


Total 
gr. wt. 

(gma.) 


Gr. wt. 
of roots 

(gms.) 


Total 

dry wt. 

(gms.) 


Greatest 

length 

(cm.) 


Initial P H 
of sol. 


1 


AcO 


25.44 


7.63 


2.325 


32.33 


3.4 


2 


Ac/10 


33.52 


7.96 


2.567 


32.91 


5.8 


3 


Ac /20 


26.14 


6.36 2.170 


30.71 


63 


4 


Ac /40 


18.28 


5.10 


1.642 


22.26 


* 


5 


AcNa 


29.94 


7.89 


2.001 


32.49 


3.45 


6 


AcM 


29 . 48 


8.10 


2.139 


27.57 


5.4 


7 


Acl/ 


31.56 


8.18 


2.337 


32.56 


3.35 


8 


Ac 3/ 


28.60 


6.06 


1.888 


32.06 


3.15 


9 


BcO 


26.46 


8.60 


2.515 


33.71 


6.6 


10 


Bel/ 


26.19 


9.34 


2.232 


33.31 


6.4 


11 


Be 2/ 


27.86 


10.29 


2.151 


33 03 


5.6 


12 


Be 5/ 


25.35 


7.75 


2.108 


30.50 


3.2 


13 


BcK 2 /10 


32.25 


11.48 


2.476 


33.51 


7.0 


14 


BcK /10 


29 . 48 


9.96 


2.453 


29.40 




15 


BcCa 


30.42 


11.80 


2.852 


34.55 


7.3 


16 


Bc/5 


31.20 


10.68 


2.530 


34.65 


7.4 


17 


Bc/10 


31.55 


9.90 


2.767 


33.98 


86 



* Heavy precipitate. 



tions, the second series was placed upon the rotating table and 
rotated throughout the period of culture. The rotating table 
employed was that previously described (Duggar and Bonns, 
'18) except that there was substituted for the special pot plat- 
forms a continuous platform constructed above the radiating 
arms and the secondary motion was, of course, eliminated. 
The wheat was grown 21 days, while the corn and peas were 
grown 24 days. One change of solution was made after about 
12 days of growth. It is, therefore, a rather severe test of 
growth quantities when infrequent renewals of solutions are 
made. The water loss from transpiration was supplied about 
every second day. 

In determining the hydrogen ion concentration of the solu- 
tions employed at the beginning of the experiment it was found 
that the active acidity of solution A was 3.4, consequently much 
greater than the theoretical. This was found to be due to the 



1920] 



DUGGAH — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



17 



TABLE VI 

(Series 2, Peas) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 



No. 



Culture 

indices 



Total 
gr. wt. 

(gms.) 



Gr. wt. 

of roots 

(gms . ) 



Total 

dry wt 

(gms.) 



Greatest 

length 

(cm.) 



P 



Initial 



After gr. 



1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 
14 
15 
16 
17 



ApO 
Ap/10 

Ap/20 
Ap/40 
ApNa 
ApM 

Apl/ 
A P 3/ 
BpO 

Bpl/ 
Bp2/ 

Bp5/ 
BpK 2 /10 

BpK /10 
BpCa 

Bp/5 
Bp/10 



16.01 
17.89 
15.34 
11.55 
20.30 
18.28 
19.39 
14.17 
15.91 
16.69 
16.58 
15.78 
16.12 
18.75 
17.15 
17.49 
18.38 



5.29 
6.21 
5.30 
4.68 
7.20 
6.15 
6.60 
5.22 
5.70 

6.13 

5.82 
5.33 
5.91 
6.87 

6.18 
6.32 

6.29 



1.351 
1.473 
1.332 
1.124 
1.603 
1.418 
1.505 
1.262 

1.326 
1.410 
1.391 
1.316 
1.413 
1.501 
1.393 
1.389 
1.413 



24.02 

22.72 
19.93 
12.49 
26.03 
24.19 
24.50 
21.13 
19.59 
22.16 
22.48 
20.24 

19.01 
21.33 
23.44 
19.21 
21.65 



3.4 

5.8 
6.3 

3.45 

5.4 

3.35 

3.15 

6.6 

6.4 

5.6 
3.2 
7.0 



7.3 
7.4 

8.6 



4.6 

5.8 
6.0 

6.8 
4.4 
6.4 
4.6 
4.2 
6.2 
6.0 
6.0 
4.6 
6.6 
5.2 

8.0 
7.4 
8.0 



* Heavy precipitate. 



use of a new supply of monobasic potassium phosphate. This 
was a high-grade reagent, but was not guaranteed free of phos- 
phoric acid, and no such guaranteed salt was then obtainable. 
The determinations indicate very clearly that attention must be 

paid to the determinations of the P H value whenever such experi- 
ments on nutrition are conducted. This series of experiments 
must be examined and interpreted in the light of the Ph value 
referred to. Nevertheless, it should be pointed out here, though 
emphasized later, that the P H exponent of the more acid solu- 
tions is rapidly increased with the growth of the crop. The 
increased acidity is doubtless due to an excess of acid in the 
preparation of the salt. This relatively high acidity affected, 

of course, to a degree the P H of other solutions to which alkali 
was added. 

In general, this series is an extension and continuation of the 
previous work, modified particularly by the addition of certain 



2 



Vol. 7 



18 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



cultures not included in the previous series. Attention may be 
drawn to the inclusion of cultures in which the hydrogen ion 
concentration of the A series was increased by slight additions of 
phosphoric acid, as in Ax 1/, Ax 3/, likewise in solution A the 
substitution for one-half the quantity of the monobasic salt by 
an equivalent of the dibasic potassium phosphate, this being 
designated AxM. In the B solution there were also introduced 
cultures to which relatively small amounts of phosphoric acid 
were added, Bx 1/, Bx 2/, and Bx 5/, also one culture in which 
was included 1 gm. of solid calcium carbonate, BxCa. A fresh 
quantity of the salt in the case last mentioned was introduced 
with each change of solution. A general examination of the 
total green weight quantities in the case of wheat indicates that 
under the conditions of this experiment the maximum growth 
quantities were obtained with the B solution. Small amounts of 
phosphoric acid or of either phosphate (BxK 2 /10 and BxK/10) 
gave an increase in growth over the unmodified solution, and 
within the range of hydrogen ion concentration which prevailed 
in this culture solution relatively little influence was exerted by 
changes in P H except in the case of one culture, Bw 5/, where 
the hydrogen ion concentration was increased to P H 3.2. On 
the other hand, with the A solution it is clear that the addition 
of alkali to the unmodified solution is generally beneficial, at 
least at concentrations up to and including Aw /10, in which 
culture the P H exponent is raised to 5.8. Further addition of 
alkali gives a falling off in the growth quantities. In the culture 
Aw /20 where the hydrogen ion concentration is P H 6.3, pre- 
cipitation occurred and a marked decline in growth is apparent, 
although the reaction of the medium is the same as that which 
in the B series promoted an amount of growth approaching the 
maximum. This is one of the many indications pointing clearly 
to the probability that the most favorable hydrogen ion con- 
centration, or range of concentration, for a particular nutrient 
solution does not necessarily correspond to that which is most 
favorable with the solution of entirely different constitution. 
The growth quantity recorded for Aw 1/ is not strictly in line 
with the discussion above. The duplicates differed consider- 
ably. We have here, of course, two factors involved: (1) 
increase in aciditv. and (2) sliehtlv increased PO4 concentration. 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 19 



With respect to corn it will be seen that the maximum yield 
occurs in culture Ac /10, although maximum root growth occurs 
in several B cultures. As between the different cultures in the 
A series the results are much the same as in the case of wheat 
except that the AcM culture with a combination of mono- and 
dibasic phosphate does not exhibit the benefit reported in the 
previous case. On the other hand, when one examines the data 
for dry weight quantities it will be found that maximum growth, 
as before, occurs in the B solution and in the culture to which 
solid calcium carbonate was added. Moreover, the next best 
growth is found in the addition of a slight amount of alkali to the 
B solution. 

Throughout this work it will be noticed that the variation in 
total growth amounts between the various cultures of peas, 
differing mainly in P H , is not so marked as that shown by the 
other two plants used in these experiments. Again, there were 
considerable differences between different plants in the same 
culture (more marked, however, in the case of wheat), and this 
indicates beyond any doubt that the variability of the seed is a 
factor which may affect to a slight extent the regularity of the 
results. In any event, the maximum growth with peas in this 
series occurred when a small amount of sodium sulphate was 
added. The amount of growth in the unmodified A solution is 
considerable in spite of the high acidity, yet there appears to be 
a slight advantage in the addition of a small quantity of alkali, 
although the latter is rendered doubtful by a comparison of 
cultures ApO, Ap 1/, and Ap 3/. In solution A Canada field 
peas are apparently only slightly affected by changes in hydro- 
gen ion concentration up to the point of precipitation of the 
phosphate as the insoluble calcium salt. The curve, fig. 2, 
exhibits all the necessary data for growth comparison. Since 
these experiments were made it has been found that a frequent 
laboratory grade of acid potassium phosphate will give a P H 
anywhere from 3.5 to 4.5, but more frequently less than 4.0. 

Change in P H after plants had grown in the solution was fol- 
lowed in the case of peas, with a result much like that of the first 
series. The more acid solutions are shifted towards neutrality, 
but solutions with exponents greater than 5.8 or 6.0 may vary 



while the more alkaline 



shifted toward 



[Vol. 7 



20 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



TABLE VII 

(Series 3, Wheat) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH; 

SOLUTIONS A, B, AND C; RENEWAL EVERY FOUR DAYS 



No. 



Culture 
indices 



Total 
gr. wt. 

(gins.) 



Total 
drywt. 

(gms.) 



Greatest 
length 

(cm.) 



P 



Initial 






i 

2 
3 
4 

5 
6 

7 

8 

9 

10 



AwO 

Aw 9/10 K 
Aw 4/5 K 
Aw 3/5 K 
Aw 1/2 K 
Aw 3/10 K 
Aw h/Al 
AwAl 

BwO 

2(BwO) 



1 1 BwK 2 



12 

13 
14 
15 
16 
17 
18 
19 
20 
21 



BwCa 
BwAl 

CwO 
CwNH 

CwGC 
CwGP 
Cw/1 

Cw/2 
Cw/5 
CwAl 



1.92 
7.92 
6.27 
6.36 

10.18 
6.55 
5.21 
6.39 
8.24 
7.65 
3.22 
6.47 

13.85 
5.60 
3.71 
5.29 
4.37 
8.44 
7.72 
6.15 
S.20 



0.325 


12.02 


1 . 054 


1804 


0.776 


17.08 


0.814 


14.82 


1.211 


20.52 


0.789 


17.12 


0.862 


17.26 


0.893 


17.60 


1.124 


19 96 


1.010 


19 . 27 


. 500 


15.00 


. 950 


21.34 


1.954 


25.33 


0.683 


14.89 


0.603 


14.87 


0.796 


16.21 


0.640 


14.10 


1.045 


18.54 


1 . 022 


16.96 


. 928 


16.23 


1.178 


20 -74 



4.8 
5.3 
5.5 
60 
6.1 
6.4 
3.6 
5.3 
5.9 
5.8 
7.1 
7.6 
71 
4.8 
4.3 
6.7 

7.1 
5.3 

5.8 
6.4 
54 



End 2nd per. 


End 4th per. 


4.8 


4.8 


5.6 


6.0 


5.5 


5.5 


5.8 


5.8 


5.7 


6.1 


61 


61 


36 


53 


5.4 


5.8 


7.1 


7.1 


7.3 


8.0 


7.4 


7.8 


7.6 


7.8 


7.4 


7.7 


5.6 


65 


4.6 


4.8 


7.0 


7.8 



7.0 
5.8 
6.0 
64 

5.8 



7.4 
7.2 

6.6 
7.3 
7.0 



Series 3 was arranged with a view to repeating some of th< 
work previously conducted and likewise to an extension of it 
Two plants, wheat and corn, were employed, and the subdivi 
sions of the series mav be appropriately designated 3w4, 3wl0 



ndicating respectively (3w4) cultures with 
red every 4 days; (3wl0) cultures with 
;ed every 10 days; (3c4) cultures with cor 



heat 



tions 



ed every 4 days; and (3c 10) cultures with corn, 



solutions changed every 10 day 



1 



this 



therefore, a 



of 



duplicated by a similar 
ved every 10 days. 



Sections 3w4 and 3< 
table, while sections 3 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



21 



TABLE VIII 

(Series 3, Wheat) 

REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GR< 
SOLUTIONS A, B, AND C; RENEWAL AFTER A TEN-DAY INTERVAL 



No. 



Culture 
indices 



Total 
gr. wt. 
(gms.) 



Total 

drywt. 

(gms.) 



Greatest 
length 

(cm.) 



P 



Initial 



1 
2 
3 
4 
5 
6 
7 

8 
9 

10 
11 
12 
13 
14 
15 
16 
17 
18 
19 
20 
21 



AwO 

Aw 9/10 K 

Aw 4/5 K 

Aw 3/5 K 
Aw 1/2 K 
Aw 3/10 K 
Aw h/Al 
AwAl 

BwO 

2(BwO) 

BwK 2 

BwCa 

BwAl 

CwO 

CwNH 

CwGC 

CwGP 

Cw/1 

Cw/2 

Cw/5 
CwAl 






2.55 
5.04 
5.50 
5.52 

5.53 
1.06 
2.52 
4.40 
7.62 
7.90 
2.82 
5.12 
10.30 
4.95 
5.57 
5.50 
1.70 
7.10 
6.42 
5.50 
6.45 






0.482 

0.765 
0.812 
0.748 
0.849 
0.262 
0.506 
0.668 
1.079 
1.094 

0.470 
0.789 
1.622 
0.707 
0.858 
0.846 
0.308 
0.995 
0.887 
0.778 
0.971 



r 



End 1st per. 



End 2nd per. 



12.40 
15.29 
14.07 
13.76 
12.23 
10.64 
13.61 
13.52 
20.12 
19.11 



4.8 
5.3 
5.5 
6.0 
6.1 
6.4 

3.6 
5.3 
5.9 

5.8 



14.42 


7.1 


17.14 


7.6 


26.65 


7.1 


14.10 


4.8 


16.15 


4.3 


16.91 


6.7 


10.77 


7.1 


16.24 


5.3 


16.62 


5.8 


16.28 


6.4 


14.67 


M 



4.6 
5.6 
5.2 

5.6 
5.5 
6.0 

3.8 
5.5 
7.6 
76 

8.2 

7.8 
7.6 
5.9 
4.6 
7.6 
7.4 
6.2 
6.4 

6.8 
6.4 



4.8 
6.3 
5.9 
6.1 
6.4 
6.1 
4.9 
6.1 
7.4 
8.0 
8.5 

8.8 
8.0 

6.7 
5.3 
8.8 
7.6 
7.7 
7.6 
7.4 
7.4 



house, and the cultures were distantly spaced on a lattice table, 
or bench, constructed with the idea of providing for all cultures 
uniform circulation, or conditions favoring uniform water loss. 
The light relations were perhaps somewhat more favorable for 



ed at 



day intervals. This 



During the first two weeks the 



the group of cultures renev 
series was begun on May 14. 
weather was moderate and cloudy, while during the last week it 
was bright and warm, perhaps too warm for the best growth of 
wheat. While the evaporation from a standardized spherical 
atmometer was low, seldom exceeding 12 gms. per day, during 
the first two weeks it rose to a maximum of 27.8 and 25.4 as a 
record for the two houses on May 31. 

The modification in the A group of cultures in this series con- 
sisted chiefly in the introduction of K 2 HP0 4 as a source of part 






22 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



TABLE IX 

(Series 3, Corn) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH; 

SOLUTIONS A, B, AND C; RENEWAL EVERY FOUR DAYS 



No. 



1 

2 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 
16 
17 
18 
19 
20 
21 
22 
23 



Culture 
indices 



Total 

gr. wt. 

(gms.) 



Gr. wt. 

of 
roots 
(gms.) 



Ac h/1 

Ach/2 

AcO 

Ac 9/10 K 

Ac 4/5 K 

Ac 3/5 K 

Ac 1/2 K 

Ac 3/10 K 

Ac h/Al 

AcAl 

BcO 

2BcO 
BcK 2 

BcCa 

BcAl 

CcO 

CcNH 

CcGC 

CcGP 

Cc/1 

Cc/2 

Cc/5 

CcAl 



9.55 

32.10 

50.61 

33.08 

41.12 

40.90 

40.80 

47.20 

16.57 

27.78 

3606 

49.72 

43.23 

3641 

37.90 

29.02 

20.44 

33.33 

32.82 

35 . 83 

35.32 

35.70 

41.83 






1.70 

7.08 
12.94 
10.24 
10.13 

9.76 
12.98 
15.24 

3.82 

7.85 

9.65 

13.75 

12.12 

11.01 

12.45 

8.58 

4.68 

9.86 

8.83 

10.82 

8.80 
10.25 
12.33 



Total 
dry 

wt. 
(gms.) 



Great- 
est 

length 
(cm.) 



1.422 
3.301 
4.716 
3.015 
3.898 
3.820 

3 . 434 
3.915 
2.117 
3.288 
4.107 
5.159 

4 . 290 
4.305 
4.597 

2 . 920 
2.307 
3.210 
3.061 

3 . 437 
3.306 
3.271 
4.201 



Ph 



Initial 



16.78 
27 . 92 
35.74 
26.58 
30.50 
29.60 

29.98 
31.75 
19.93 
23 . 05 
29.32 
34.11 
30.94 

29.78 
32.46 
23.95 
22.44 
25 . 82 
26.57 
2!). 01 
27.84 
28.18 
29 . OS 



3.4 

3.7 
4.8 
5.3 
5.5 

6.0 

6.1 

6.4 

3.6 

5.3 

5.9 

5.8 
7.1 

7.6 

7.1 

4.8 
4.3 
6.7 
7.1 

5.3 

5.8 
6.4 
5.4 



End 2nd per. 



End 4th per. 






3.4 
4.1 
4.8 
5.6 
53 
5.6 
5.6 
6.0 
4.6 
5.2 
6.8 
7.1 
7.0 
7.2 
7.1 
5.2 
4.6 
6.8 
6.6 
5.6 
6.1 
6.4 
57 



5.2 

5.6 

5.6 

5.8 
6.2 

6.3 
6.9 
6.9 
53 
6.3 
6.8 
7.0 
7.0 
7.2 
5.7 
7.2 
4.6 
7.7 

7.8 
7.8 
7.6 
8.0 
7.6 



of the potassium and phosphate ions. The stock solution of 
this salt was made to contain the same number of gram atoms 
of P0 4 as the stock solution of KH2PO4. These solutions were 
then combined so that the ratios of monobasic to dibasic phos- 
phate were respectively 9 : 1,4: 1,3: 2, 1 : 1, and 3: 7, the culture 
indices being Ax 9/10 K, Ax 4/5 K, Ax 3/5 K, Ax 1/2 K, and 
Ax 3/10 K, with the P H of the solutions affected as indicated in 
the tables. In culture AxO, another high grade of KH 2 P0 4 was 
employed, and the P H exponent is higher than the theoretical; 
while in the Ax h/1 cultures, used especially in the case of corn, 
the more acid grade is employed. Ac h/2 is intermediate in 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



23 











TABLE X 










(Series 3, Corn) 






SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION ' 


TO GROWTH; 


SOLUTIONS A, B 


i. AND C; RENEWAL AFTER A TEN-DAY INTERVAL 




1 


m t 1 


Gr. wt. 


Total 


Great- 




Ph 






Culture 
indices 


Total 


of 


dry 
wt. 


est 1 








No. 


gr. wt. 


roots 


length 








' 


(gms.) 


(gms.) 


(gms.) 


(cm.) 


Initial 


End 1st per. 


End 2nd per. 


1 


Ach/l 


13.70 


3.15 


2.080 


22.42 


3.4 


5.1 


6.4 


2 


Ach/2 


18.13 


3.83 


2.268 


23.77 


3.7 


5.4 


6.3 


3 


AcO 


34.42 


7.75 


3.922 


31.65 


4.8 


4.0 


5.2 


4 


Ac 9/10 K 


34.82 


9.10 


3.282 


2964 


5.3 


5.6 


5.3 


5 


Ac 4/5 K 


1 34.12 


7.72 


3.298 


27.17 


5.5 


5.1 


5.3 


6 


Ac 3/5 K 


37.62 


9.70 


3.589 


28.07 


60 


56 


54 


7 


Ac 1/2 K 


41.80 


11.30 


3.784 


28.76 


61 


5.6 


5.4 


8 


Ac 3/10 K 


37.50 


12.56 


3.397 


27.25 


6.4 


5.8 


6.4 


9 


Ac h/Al 


19.67 


4.34 


2.491 


23.23 


3.6 


5.2 


6.1 


10 


AcAl 


32.29 


9.22 


3.389 


25.70 


5.3 


5.8 


5 3 


11 


BcO 


28.26 


8.78 


3.366 


24.36 


5.9 


7.2 


5.5 


12 


2BcO 


3804 


7.77 


4.525 


30.22 


5.8 


7.7 


5.3 


13 


BcK 2 


43.00 


12.63| 4.172 


31.83 


7.1 


8.0 


5.8 


14 


BcCa 


29.85 


9.20 


3.617 


26.59 


7.6 


8.4 


7.3 


15 BcAl 


27.80 


7.70 


3.271 


27.75 


7.1 


7.0 


5.6 


16 


CcO 


30.65 


8.40 


2.923 


24.84 


4.8 


5.6 


7.6 


17 


CcNH 


14.44 


3.80 


1.994 


23.85 


4.3 


4.6 


5.8 


18 


CcGC 


27.50 


6.97 


2.788 


23.94 


6.7 


7.6 


8.4 


19 


CcGP 


32.37 


8.90 


3.145 


23.83 


7.1 


7.3 


8.2 


20 


Cc/1 


31.85 


7.20 


3.060 


26.95 


5.3 


6.2 


7.5 


21 


Cc/2 


34.50 


9.35 


3.005 


29.65 


5.8 


64 


7.4 


22 


Cc/5 


38.90 


9.25 


3.487 


29.43 


6.4 


7.0 


7.1 


23 


CcAl 


25.06 


5.89 


i 2.699 


24.74 


5.4 


6.4 


7.7 



acidity between AcO and Ac h/1, with salt proportions the same. 
Again, to other cultures containing these two grades of phos- 
phates there were added to each culture vessel (and with each 
renewal) 1 gm. of solid aluminium hydroxide of the highest 



purity procurable 
Al respectively. 



designated AxAl and Ax h/1 



In the B group cultures were prepared with the addit 



solid 



uminium hydroxide (BxAl), as abo 



also 



ith solid 



arbonate BxCa, with the addition of a small amount of 



K2HPO4 (BxK a ), and the unmodified solution of double 
[2(BxO)l. 



oa [VoL - 7 

^4 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



The C group of cultures (Livingston-Tottingham medium) is 
here introduced for the first time. Explanations already given 
explain the culture indices in this group, except in the following 
cases: CxNH, CxGC, and CxGP. In the first mentioned one- 
half the atomic proportion of nitrogen is supplied as (NH 4 ) 2 S0 4 , 
and in the last two calcium glycero-phosphate is added to re- 
place monobasic calcium phosphate. In CxGC the atomic 
proportion of Ca is kept the same as in CxO, while in CxGP it 
is the phosphorus which is equivalent. 

Under the conditions of these experiments the Aw cultures, 
particularly, were less satisfactory than usual, possibly in large 
part due to the high temperature prevailing towards the end of 
the period. The series was discontinued earlier than planned 
owing to the drying of the leaf tips of wheat and even of some 
entire plants. On the whole the wheat cultures show many 



results. 



but despite this also some strikin 



Increasing the P H exponent by means of the dibasic phosphate 
may under these extreme conditions more than treble or quad- 
ruple the growth quantities. It is probably a matter of shifting 
the sum of conditions from the side of toxicity to that of growth 
maintenance. Even in the case of the C solution, increasing the 
P H exponent is here a factor in promoting growth increase. 
Under other conditions I have not found this to be true, as will 
be indicated later. 

The addition of aluminium hydroxide is under these condi- 
tions distinctly favorable, as seen by comparing the following 
pairs, AwO and AwAl (also AwO and Aw h/Al), BwO and 
BwAl, CwO and CwAl. The value of this reagent is doubtless 
in part due to its action as a buffer. 

The temperature was most favorable for the corn cultures, 
and they exhibited, on the whole, an unusually vigorous growth. 
The green weight determinations are an accurate indication of 



growth extent but 



dark 



green in color, with heavy purplish stems ; while both the A and 
C groups were strongly chlorotic. Chlorosis was much intensi- 
fied during the last week of growth, and it seemed probable 
that if these cultures were longer maintained, a considerable 
reduction in the growth rate would occur. It is assumed, how- 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 25 




ft 



o^^^^^^^^e^-^r^^^ja/fi^ census jb^ju^a^ 



V.3 



CJftCLAP Cur't 
6.7 77 J-3 



60 

Cu/"£ 



6/ 

Cur4 



6.+ $6 



53 SS S-S 



7< 



7* 7A 



Fig. 3. Total green weight of wheat in solutions A, B, and C, with modifications 
effected largely by combinations of phosphates or by the introduction of "insoluble" 
buffers. Intervals between the renewal of solutions were 4 and 10 days. 



ever, that in these cases the chlorosis may be related to inade- 
quate iron supply, and not to faulty proportions of the main salt 
constituents. 

The Ac h/1, Ac h/2 and Ac h/Al cultures all show the in- 
jurious effect of high acidity; but with the initial value of P H 4.8 
the best growth in the one lot is in the A solution, though practi- 
cally approached in the double strength of the B solution 
[2(BcO)]. Aside from the considerable variation in cultures 
differing only slightly in composition or in P H , the chief point 
of interest is the depressing action of the ammonium salt in both 
lots (BcNH). 

The curves, figs. 3 (wheat) and 4 (corn), exhibit diagrammat- 
ically the data above discussed, and require no explanation, 
further than to point out that the results with solution C appear 



[Vol. 7 



26 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



(Pom v , 1% 

Soi.C(¥Ja) 
Sal.CC'Od*) 





2k*tltaiL&i&.d*>t£Mi4*h\ 



BffH (mi. ..$*MuA«»A. S.&flfe«Jl 



cu k / cutt. cucr cu%^ac%^ac%KOcy 2 Kac%ic(Aad2i Bco-^co jbcTk ag B*ai 



7i/ 5* 57 V8 ' $3 $5 6.0 6J 6.4- 

J CdCf CcW Cc5C CctfP Cc4 Cc'z Cc y 5 t CcQt 

<h VS y 3 6-7 7/ y.3 jr.3 6¥ **■¥ 



3,6 



S3 S.9 5.5 J.i 



7* 7 



./ 



Fig. 4. One-fourth total green weight of corn in solutions A, B, and C. See 
further explanation under fig. 3. 



the same reeion of fig, 3 as solution A merely because 



necessarj 
figure 



do this in order to present all the dat 



order to include solution C again in the tests, and 
same time to change somewhat the range of hydrog 



ion 



concentrations 



and B solutions, as well as to repeat 



the former work, a more 



ments was 



ed with wheat, corn, and peas as indicated by 



in 



tables xi-xix. Meanwhile, it had been determined from 
liminary experiments that nutrient solutions of diverse con- 
ution seem to be considerably influenced by the conditions 

cultures were grown. There was, therefore, 



under which the cultures were grown. 

introduced into this series three sets of condit 

of the fact that other experiments were under way in the green- 



On 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



27 



TABLE XI 
(Series 4, I — Wheat [moist, high temperature]) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 



No. 



Culture 
indices 



Total 
gr. wt. 
(gms.) 



Total 
dry wt 

(gms.) 



Greatest 

length 

(cm.) 



P 



H 



Initial 



After gr. 



1 
2 

3 
4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

21 



AwO 
Aw /5 
Aw /10 
Aw 1/ 
Aw 3/ 
AwNa 
AwM 
BwO 
Bw /5 
Bw/10 
Bw 1/ 

Bw5/ 

Bw 10/ 

BwK 2 /10 

BwCa 

CwO 

Cw/5 

Cw/20 

Cw 1/ 

Cw5/ 

CwNa 



4.19 


.491 


30.71 


3.4 


4.10 


.400 


26.77 


4.6 


3.78 


.399 


27.00 


5.8 


2.60 


.338 


24.63 


3.35 


2.22 


.328 


22.08 


3.15 


3.83 


.468 


30.10 


3.45 


4.65 


.485 


29.23 


5.4 


5.22 


.469 


32.96 


6.6 


4.90 


.466 


32.84 


7.4 


4.65 


.424 


32.04 


8.6 


5.38 


.491 


32.06 


6.4 


5.35 


.472 


31.33 


3.2 


.95 


.177 


12.26 


2.8 


5.13 


.466 


35.55 


7.0 


5.70 


.539 


34.23 


7.3 


4.82 


.465 


29.12 


4.2 


4.17 


.401 


24.78 


6.4 


2.25 


.301 


18.85 


7.6 


4.05 


.403 


26.50 


3.6 


2.47 


.296 


20.85 


3.2 


4.50 


.441 


28.72 


4.2 



4.0 

5.3 
4.4 
4.4 
3.6 
3.8 
4.0 
7.2 
7.6 
8.2 
7.2 
6.8 
3.8 
7.2 
6.6 
6.4 
7.0 
7.2 
6.4 
5.2 
6.6 



houses at the same time no particular effort was made to con- 
trol accurately the conditions of growth. The following general 
conditions were decided upon: (i) Moist high temperature, (n) 
moist low temperature, and (in) dry high temperature. These 
conditions are relative, of course, and all supported good growth. 
"Moist" in the sense here used simply means that by frequent 
sprinkling of walls and floors the humidity was raised above that 
of the usual greenhouse compartment. However, in the case 
of those cultures placed under conditions of high moisture and 
high temperature there was also necessary a slight degree of 
shade. This caused the plants to grow up rather quickly. As 
a result of this rapid growth in this section of the series it was 
determined to take down all cultures after about 15 days except 
those placed under the conditions referred to as moist, low 



28 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



TABLE XII 

(Series 4, II — Wheat [moist, low temperature]) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 



^ T 


Culture 


Total 

■ 


Total 


Greatest 


Ph 


No. 


^-^ V4A V VAX \> 

indices 


gr. wt. 


dry wt. 


length 








(gms.) 


(gms.) 


(cm.) 


Initial 


After gr. 


1 


AwO 


3.72 


.681 


16.81 


3.4 


4.0 


2 


Aw /5 


10.87 


1.373 


22.90 


4.6 


6.0 


3 


Aw/10 


8.85 


1.087 


21.40 


5.8 


6.4 


4 


Aw 1/ 


4.42 


.736 


16.75 


3.35 


4.0 


5 


Aw 3/ 


3.27 


.579 


16.23 


3.15 


3.4-3.5 


6 


AwNa 


6.80 


.862 


19.28 


3.45 


5.8 


7 


AwM 


11.24 


1.323 


24.20 


5.4 


6.6 


8 


BwO 


10.92 


1.035 


20.75 


6.6 


7.0 


9 


Bw/5 


8.40 


.912 


19.27 


7.4 


7.6 


10 


Bw/10 


9.02 


.924 


22.12 


8.6 


8.1 


11 


Bw 1/ 


7.37 


.908 


20.69 


6.4 


7.1 


12 


Bw5/ 


9.17 


.910 


20.06 


3.2 


7.2 


13 


BwlO/ 


.90 


.209 


8.80 


2.8 


3.6 


14 


BwK 2 /10 


8.82 


.984 


19.46 


7.0 


7.5 


15 


BwCa 


11.64 


1.354 


20.51 


7.3 


7.4 


16 


CwO 


6.75 


.870 


18.54 


4.2 


7.7 


17 


Cw/5 


6.77 


.826 


16.93 


6.4 


7.8 


18 


Cw /10 


5.60 


.681 


12.85 


7.6 


8.6 


19 


Cw 1/ 


7.22 


.915 


18.90 


3.6 


7.7 


20 


Cw5/ 


7.55 


.916 


18.17 


3.2 


7.5 


21 


CwNa 


7.75 


.853 


18.78 


4.2 


7.8 



temperature. On account of conditions prevailing at the time, 
sections i and in were actually maintained from 16 to 18 days. 
The plants in section n were permitted to run from 25 to 27 



days 



This was an extreme test of the tolerance of these 
and no change of the nutrient medium was made ii 



culture throughout the entire 



d of erowth. The water 



by transpiration was, however, replaced from day to day 



A general survey of 



blocks of cultures represented by 



without chan 



ns A, B, and C gives evidence that under conditions of 
moist air and high temperature (table xi) wheat grown 

;e of solution for 18 days exhibits less striking 



the maximum growth quantities than may be 



of 



Nevertheless, the 



of total green weight or of dry weight, is found 



1920] 






DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



29 



TABLE XIII 

(Series 4, III — Wheat [dry, high temperature]) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 



No 



Culture 
indices 



Total 
gr. wt. 
(gms.) 



Total 
dry wt. 

(gms.) 



Greatest 
length 

(cm.) 



P 



H 



Initial 



After gr 



1 
2 
3 
4 
5 
6 
7 

8 
9 
10 
11 
12 
13 
14 

15 
16 
17 

18 
19 
20 
21 



AwO 

Aw /5 
Aw /10 
Aw 1/ 
Aw 3/ 
AwNa 
AwM 

BwO 

Bw/5 

Bw/10 

Bw 1/ 
Bw5/ 
Bw 10/ 

BwKs/10 

BwCa 

CwO 

Cw/5 

Cw/20 

Cwl/ 

Cw5/ 
CwNa 



2.77 
5.50 
5.61 
1.78 
1.55 
3.66 
3.35 
7.43 
7.25 
564 
4.72 
5.42 
.17 
6.40 
7.39 
4.49 
4.97 
3.32 
4.80 
2.00 
4.50 



.395 

.582 
.601 
.287 
.248 
.441 
.390 

.658 
.733 
.493 
.554 
.566 
.092 
.615 
.765 
.510 
.576 
.450 
.491 
.298 
.522 



19.70 
26.23 
24.90 
17.14 
16.33 
23.70 
21.89 
2654 
30.97 
22.37 
24.38 
25.19 
6.67 
26.58 
28.62 
21.60 
23.43 
20.20 
23.66 
19.02 
24.00 



3.4 

4.6 
5.8 
3.35 
3.15 

3.45 

5.4 

66 

7.4 
8.6 
6.4 
3.2 

2.8 
7.0 
7.3 
4.2 
6.4 
7.6 
3.6 
3.2 
4.2 



4.8 
5.8 
6.1 
3.4 
3.4 

3.6 
5.0 
7.2 
7.6 
8.6 
8.6 
7.3 
2.9 
7.9 
8.2 
7.2 
7.6 
8.2 
7.0 
5.0 
7.4 




BwCa. Several other cultures in the B block, notably Bw 1 
and Bw 3/, exceed slightly all those of the C block by either of 
the 2 important criteria, that is, green weight or dry weight. 
Under these conditions the tolerance of high hydrogen ion con- 
centration is generally marked, as shown in cultures AwO, 
AwNa, Bw 5/, and Cw 1/. Likewise the range of tolerance 
and of strong growth is considerable. 

In section n of this series with wheat (table xn) it is the infer- 
ence from the data that when grown for a longer period (25 
days) under cooler conditions wheat is less resistant to high 
hydrogen ion concentration. This is shown in part by the fact 
that the 4 cultures yielding highest in green weight are BwCa, 
AwM, BwO, and Aw /5. There is, however, no striking falling 
off, on the whole, as hydrogen ion concentration increases to 



30 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



TABLE XIV 



(Series 4, I 



temperature]) 



SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 



No. 



1 

2 
3 
4 
5 
6 

7 
8 
9 
10 
11 
12 
13 
14 
15 
16 
17 
18 
19 
20 
21 



Culture 
indices 



AcO 

Ac/5 

Ac/10 

Acl/ 

Ac 3/ 

AcNa 

AcM 

BcO 

Bc/5 

Bc/10 

Bel/ 

Be 5/ 

Be 10/ 

BcK 2 /10 

BcCa 
CcO 
Cc/5 
Cc /20 

Cc 1/ 
Cc5/ 
CcNa 



Total 
gr. wt. 
(gins.) 



Gr. wt. 

of roots 

(gms.) 



Total 

dry wt 
(gms.) 



Greatest 
length 

(cm.) 



P 



H 



Initial 



19.05 


4.42 


.593 


34.72 


27.32 


6.80 


1.786 


40.20 


18.85 


3.87 


.305 


38.58 


14.83 


4.01 


1.136 


30.48 


9.45 


3.28 


1.129 


29.23 


6.44 


3.09 


.973 


22.77 


24.02 


4.80 


1.615 


38.24 


22.31 


4.93 


1.781 


38.63 


19.77 


2.44 


1.452 


39. 65 


21.12 


4.97 


1.538 


39.64 


20.93 


3.81 


1.726 


40.85 


21.55 


4.33 


1.724 


41.25 


9.02 


2.38 


.936 


22.71 


25.56 


4.44 


1.770 


43.46 


19.02 


2.70 


1.572 


43.40 


20.02 


4.70 


1.396 


27.26 


16.11 


3 . 28 


1.105 


26.72 


15.81 


4.81 


1.207 


22.18 


19.81 


4.88 


1.341 


33.56 


26.72 


6.67 


1.780 


31.02 


18.81 


3.94 


1.165 


30.62 



3.4 

4.6 

5.8 

3.35 

3.15 

3.45 

5.4 

6.6 
7.4 
8.6 
6.4 
3.2 
2.8 
7.0 
7.3 
4.2 
6.4 
7.6 
3.6 
3.2 
4.2 



After gr 



4.2 
4.6 
6.0 
6.3 
6.4 

6.6 
65 
5.2 
6.4 
6.1 
5.6 
4.4 

7.1 
5.2 

7.2 

7.7 
7.4 
7.9 
7.6 
7.6 
7.7 



about P H 3.2, although there is not the consistency which might 
be expected between cultures AwO, Cw 1/, Cw 5/, and Bw 5/. 

Where the conditions involved a relatively dry atmosphere 
and a high greenhouse temperature the response of the organism 
to the different culture solutions is of special interest. The 



block, nam 



with highest green weight yields are all in the 



BwO, BwCa. and 



/5. These are so far 



ahead of the cultures in the A and C blocks, irrespective of 
hydrogen ion concentration within the usual range, as to leave 
no doubt whatever that for continued cultivation in the same 
solution for a period of 18 days the B solution is decidedly the 
most favorable culture medium. In this case too we have as 
definite an indication as has been afforded as to the limitation 
imposed by hydrogen ion concentrations. With an exponent 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



31 



TABLE XV 

(Series 4, II — Corn [moist, low temperature]) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 



No 



Culture 

indices 



Total 
gr. wt. 

(gms.) 



Gr. wt. 

of roots 

(gms.) 



Total 
dry wt. 

(gms.) 



Greatest 

length 

(cm.) 



P: 



Initial 



After gr. 



1 

2 
3 
4 
5 
6 
7 
8 
9 
10 
11 



AcO 

Ac/5 

Ac/10 

Ac 1/ 

Ac 3/ 

AcNa 

AcM 

BcO 

Bc/5 

Bc/10 

Bel/ 



12 


Be 5/ 


13 


Be 10/ 


14 


BcK 2 /10 


15 


BcCa 


16 


CcO 


17 


Cc/5 


18 


Cc/20 


19 


Ccl/ 


20 


Cc5/ 


21 


CcNa 



8.77 
22.81 
23.10 
21.97 
11.28 
19.30 
27.56 
18.90 
31.99 
28.63 
23.61 
27.80 
15.50 
36.30 
26.03 
34.60 
28.47 
20.90 
23.52 
21.22 
24.60 



3.25 
8.44 
8.94 
6.94 
2.82 
7.91 

10.69 
7.50 

14.50 

14.54 
7.74 

10.60 
6.20 
6.44 

10.11 
11.02 

9.72 
6.75 
7.55 
6.70 
9.07 



1.007 
1.703 
1.521 
1.933 
1.303 
1.744 
2.078 
1.828 
2.508 
2.213 
2.423 
2.464 
1.366 
2.950 
2.699 
2.763 
2.350 
1.868 
1.966 
1.687 
2.090 






15.83 

22.98 

22.78 

21.75 

15.65 

20.56 
22.72 

20.64 

23.51 
21.84 
24.14 
23.44 
16.19 
23.50 
23.57 
24.66 
23.59 
20.10 
23.40 
22.15 
19.74 



3.4 

4.6 

5.8 

3.35 

3.15 

3.45 

5.4 

6.6 

7.4 

8.6 
6.4 

3.2 

2.8 

7.0 

7.3 

4.2 

6.4 

7.6 

3.6 

3.2 

4.2 



5.6 
6.0 
6.2 
5.8 
6.0 
6.0 
6.0 
6.4 
5.9 
7.0 
4.8 
6.0 
5.6 
5.7 
7.4 

7.6 
7.6 
8.6 

7.6 

7.6 
7.6 



below P H 4 there is definite diminution of growth in every in- 
stance, except Bw 5/. AwM seems to be distinctly out of 



harmony 



from culture Bw 710 



that under these climatic conditions a relatively low degree of 
alkalinity, probably about P H 8, would represent the limit for 
most favorable growth in a solution thus constituted. 

The results with corn grown in a moist atmosphere at a high 
temperature are perhaps more erratic than in any other section 
of the work. At least with the data at hand it is extremely 
difficult to interpret these 
best growth is in culture 



would 



Ac /5, and it is 



followed by 



Cc 5/, the last solution being close to the usual limit of growth 
respect to hydrogen ion concentration. In both these cases t 
increase in the growth over that in BcK 2 /10, AcM, and BcO 



32 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



TABLE XVI 
(Series 4, III — Corn [dry, high temperature]) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 



No. 



1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 
14 
15 
16 
17 
18 
19 
20 
21 



Culture 
indices 



Total 
gr. wt. 
(gms.) 



Gr. wt. 

of roots 

(gms.) 



Total 
dry wt. 

(gms.) 



AcO 

Ac/5 
Ac /10 
Ac 1/ 
Ac 3/ 
AcNa 
AcM 
BcO 
Bc/5 

Be /10 

Be 1/ 

Be 5/ 
Be 10/ 
BcK 2 /10 
BcCa 
CcO 
Cc/5 
Cc/20 
Ccl/ 

Cc5/ 

CcNa 



24.43 
25.90 
17.22 
21.80 
17.49 
16.00 
30.41 
19.89 
20.89 
20.90 
23.00 
20.02 

1.64 
24.52 
23.44 
16.62 
18.52 
11.28 
20.56 

8.63 
1554 






8.59 
7.94 
5.28 
5.60 
4.61 
4.70 
9.61 
6.30 
7.11 
5.85 
9.25 
7.70 
.43 
9.89 
8.81 
4.97 
5.20 
4.06 
5.65 
1.61 
5.57 



1.812 

1.751 
1.360 
1.667 
1.405 
1.231 
2.014 

1.604 
1.756 
1.810 
1.637 
1.442 

.285 
1.665 
1.867 
1.125 
1.180 

.835 
1.407 

.856 
1.122 



Greatest 

length 

(cm.) 



24.35 
22.81 
27.64 
31.24 
23.65 
27.17 
32.26 
27.07 
29.80 
30.41 
27.75 
29.59 
5.11 
31.33 
32.53 
25.67 
32.71 
14.88 
29.97 
20.75 
25.31 



P 



Initial 






3.4 
4.6 

5.8 

3.35 

3.15 

3.45 

5.4 

6.6 

7.4 

8.6 

6.4 

3.2 

2.8 

7.0 

7.3 

4.2 

6.4 

7.6 

3.6 

3.2 

4.2 



After gr 



5.9 
6.0 
6.2 
4.0 
4.6 
5.6 
6.2 

4.0 
6.8 
6.8 
5.0 
4.2 
3.4 
5.2 
7.8 
7.8 
8.0 
8.0 
7.9 
7.6 
7.8 



largely an increase in the growth of roots. It is perhaps possible 
that the partial shade of this series referred to above has been a 
factor in the irregularities which prevail throughout. There is 
no basis on which to explain the growth in AcNa. Turning, 
however, to table xv, indicating the results with corn grown at 
low temperature and higher humidity the data are notably 
different from the preceding. In this case the most favorable 
medium is culture BcK 2 /10, rather closely followed by CcO and 
less closely by Bc5/. In this section it is also notable that the 
maximum root growth occurs over a range of hydrogen ion 
concentration from 4.2 to 8.6. Exclusive of culture CcO the 
higher growth quantities are obtained for the different solutions 
at relatively low hydrogen ion concentration, that is, with media 
approaching neutrality more or less. This is particularly ob- 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



33 



TABLE XVII 
(Series 4, I — Peas [moist, high temperature]) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 



%T 


Culture 


Total 


Gr. wt. 

1* 1 


Total 

1 | 


Greatest 


Ph 


No. 


indices 


gr. wt. 


of roota 


dry wt. 


length 








(gms.) 


(gms.) 


(gms.) 


(cm.) 


Initial 


After gr. 


1 


ApO 


12.53 


4.15 


1.010 


28.46 


3.4 


3.5 


2 


Ap/5 


13.82 


4.15 


1.059 


31.05 


4.6 


4.2 


3 


Ap/10 


13.92 


3.87 


1.062 


13.25 


5.8 


5.8 


4 


Apl/ 


11.55 


3.42 


.937 


28.00 


3.35 


4.7 


5 


Ap3/ 


11.93 


3.17 


.990 


29.68 


3.15 


5.8 


6 


ApNa 


12.56 


3.93 


1.024 


31.42 


3.45 


4.6 


7 


ApM 


12.81 


3.93 


.966 


30.00 


5.4 


4.6 


8 


BpO 


10.60 


3.02 


.839 


25.87 


6.6 


6.4 


9 


Bp/5 


13.17 


3.64 


.957 


31.97 


7.4 

- 


7.5 


10 


Bp/10 


9.80 


3.11 


.746 


32.47 


8.6 


7.6 


11 


Bpl/ 


11.17 


3.47 


.809 


23.37 


6.4 


7.0 


12 


Bp5/ 


11.12 


2.89 


.842 


26.22 


3.2 


5.8 


13 


BplO/ 


7.59 


2.21 


.623 


23.29 


2.8 


4.8 


14 


BpK 2 /10 


12.30 


3.61 


.936 


26.62 


7.0 


7.3 


15 


BpCa 


12.55 


3.50 


.937 


29.57 


73 


7.6 


16 


CpO 


12.44 


3.39 


.974 


25.34 


4.2 


5.2 


17 


Cp/5 


8.19 


2.16 


.657 


21.66 


6.4 


6.2 


18 


Cp/20 


10.52 


2.93 


.928 


22.37 


7.6 


7.5 


19 


Cpl/ 


12.50 


3.50 


.982 


26.02 


3.6 


5.0 


20 


Cp5/ 


6.93 


2.01 


.605 


21.36 


3.2 


4.2 


21 


CpNa 


10.72 


3.06 


.923 


22 . 35 


4.2 


5.4 



servable in cultures Ac /10, Be /5, Be /10, BcCa, and Cc /5. 
Corn grown under conditions of high temperature and low 
humidity exhibits a maximum in AcM followed respectively by 
BcK 2 /10, Ac /5, BcCa, and Be 1/. An examination of block A 
would seem to indicate that P H 3.4 to 4.6 is entirely favorable in 
this medium, but there is a striking difference between Ac/ 10 
and AcM, which is not readily explainable* 

In table xvn it may be noted again that slight variations in 
the culture medium do not materially affect the growth of peas 
when grown under the conditions there indicated. The A 
solution with the addition of alkali yields, it is true, the maxi- 
mum growth quantities, but these quantities are only slightly in 
excess of those obtained with the same solution unmodified or 
of several cultures in the B block, notably Bp /5, BpK 2 /10, and 

3 



34 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



TABLE XVIII 
(Series 4, II — Peas [moist, low temperature]) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 



No 



Culture 
indices 



Total 
gr. wt. 
(gms.) 



Gr. wt. 

of roots 

(gms.) 



Total 

dry wt 
(gms.) 



Greatest 

length 

(cm.) 



P 



Initial 



After gr. 



1 

2 
3 
4 

5 
6 

7 

8 
9 
10 
11 
12 
13 
14 
15 
16 
17 
18 
19 
20 
21 



ApO 

Ap/5 

Ap/10 

Apl/ 
Ap3/ 
ApNa 
ApM 

BpO 

Bp/5 
Bp/10 

Bpl/ 
Bp5/ 

BplO/ 

BpK 2 /10 

BpCa 

CpO 

Cp/5 

Cp/20 

Cpl/ 

Cp5/ 

CpNa 



17.72 
25.71 
24.26 
21.05 
22.52 
20.69 
25.65 
13.87 
13.65 
18.57 
1693 
16.14 
9.55 
1811 
20.17 
16.44 
16.31 
12.32 
16.36 
17.62 
16.54 






6.55 
12.17 
9.72 
9.00 
9.44 
8.52 
11.20 
8.26 
5.18 
7.54 
6.61 
7.59 
3.88 
8.90 
8.23 
644 
6.10 
4.54 
6.95 
7.06 
5.62 



1.772 
2.290 
2.299 
1.980 
2.183 

.889 
2.310 
1.400 
1.308 
1.704 
1.590 
1.378 

.881 
1.455 
1.944 
1.650 
1.762 
1.406 
1.599 
1.760 
1.711 



19.07 
18.06 
20.00 
17.77 
19.18 
17.61 
19.87 
15.82 
14.74 
15.63 
17.38 
13.30 

10.98 
16.26 

19.28 
13.85 
13.62 
10.82 
13.61 
13.22 

17.38 



3.4 

4.6 

5.8 

3.35 

3.15 

3.45 

5.4 

6.6 

7.4 

8.6 

6.4 

3.2 

2.8 

7.0 

7.3 

4.2 

6.4 

7.6 

3.6 

3.2 

4.2 



5.5-5.6 

6.0 

5.9-6.0 

5.9 

5.6 

5.7 



6.1-6.2 

7.1 

7.6 

6.0 

6.0 

5.5 

6.2 

7.4 

7.2 

7.8 
7.7 
7.2 
6.4 
7.0 



BpCa, as also by two cultures in the C block, namely CpO and 
Cpl/. 

Wider differences are found in the case of peas grown at lower 
temperature in more humid air, but the maximum growth occurs 
in the A block, especially with the addition of a small amount of 
alkali (Ap /5) and in the ApM culture, containing both mono- 
basic and dibasic phosphates. No culture in the B block 
approaches the values referred to, and the same is true of the C 
cultures. Grown 16 days under dry conditions at a high tem- 
perature the yield of the various cultures is much reduced, and 
the relative values of the culture media do not remain the same 
as before. In this instance the B block exhibits the highest 
yields, especially cultures BpCa and BpK 2 /10. The next higher 
yield is found in Ap /10. In the A block the effect of high 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



35 



TABLE XIX 
(Series 4, III — Peas [dry, high temperature]) 

SALT REQUIREMENTS AND H-ION CONCENTRATION IN RELATION TO GROWTH 



No. 



Culture 
indices 



Total 
gr. wt. 

(gms.) 



Gr. wt. 

of roots 

(gms.) 



Total 
dry wt. 

(gms.) 



Greatest 
length 

(cm.) 



P 



Initial 



After gr. 



1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 



ApO 

Ap/5 

Ap/10 

Apl/ 

A P 3/ 

ApNa 

ApM 

BpO 

Bp/5 

Bp/10 

Bpl/ 
Bp5/ 

BplO/ 

BpK 2 /10 

BpCa 



16 


CpO 


17 


Cp/5 


18 


Cp/20 


19 


Cpl/ 


20 


C P 5/ 


21 


CpNa 



7.42 
10.62 
13.95 

3.20 

6.78 
11.22 
10.00 
12.79 
12.97 
12.90 

8.64 
11.95 

2.97 
14.32 
14.44 

6.60 
11.65 

5.11 
10.97 

5.40 

8.75 




3.05 

3. 

4.49 

1.10 

2.47 

3.60 

3.89 

4.37 

4.15 

4.12 

2.15 

4.27 

0.75 

4.90 

4.92 

2.10 

3.50 

1.22 

3.45 

1.25 

2.45 



.952 

1.026 

1.161 

.642 

.796 

.945 

1.095 

1.023 

1.059 

1.027 

.877 

.948 
.432 

1.130 

1.233 

.929 

1.046 

.760 

.977 

.636 

.966 



23.68 
24.63 
27.63 
11.82 
18.84 
25.33 
23.57 
24.06 
25.96 
26.05 
21.11 
24.23 
9.95 
27.37 
26.33 
1691 
21.43 
15.40 
21.28 
15.54 
18.99 



3.4 

4.6 

5.8 

3.35 

3.15 

3.45 

5.4 

6.6 

7.4 

8.6 

6.4 

3.2 

2.8 

7.0 

7.3 

4.2 

6.4 

7.6 

3.6 

3.2 

4.2 



4.6 
5.0 
5.4 
53 
5.0 
4.4 
5.6 

4.5 
7.3 
7.4 
7.0 
6.1 
3.0 
7.3 
7.8 
6.0 
6.6 

7.6 

5.4 

5.8 
6.0 



hydrogen ion concentration is marked. In general, as the con- 
ditions of evaporation are intensified it would appear that some- 
what more favorable results are obtained with this plant when 
the hydrogen ion concentration does not approach too closely 

the acid limit. 

It is clear, however, that the preparation of nutrient solutions 



acid phosphate 



though of high grade — may mean 



and often does mean, a hydrogen ion concentration either peril- 
ously near the critical region for growth, or actually inhibiting 
growth. This is particularly true for wheat, and it may be true 
for corn and other crops under any conditions which may accen- 

injury. Pronounced diminution in growth may occur 

as the P H exponent is progressively diminished from about 4.5. 

For each plant in the preceding series a separate set of curves 

(figs. 5, 6, and 7) has been prepared; and in each figure there are 



acid 



36 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 




o 



T5 
O 

- 



a; 

t3 



03 



a; 

a 



T3 

a 



2 

> 



o 

PQ 




2 ^ 

8 SI 






O-g 



53 .2 

bO 



G 

o 



o ' 



. 03 

CO 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 



37 







c 



03 

03 

E 



c3 

03 

s 

c 

C 

CO 

C 



.2 

> 

p£3 



C 

PQ 

GO 

O 







O ; 

GO 

03 

s 



03 



s 

03 



o 



e8 T3 



03 

o 



, o 



<D 



O 



38 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 




a 

o 

GO 

cs 
o 



> 



o 

g 

CO 

d 

O • 

CO 

> 

o 



2 
3 



3 



43 



C3 
O 



fl o 
c3 o 



o 



a 



0> 



bC o 

Bb to 

— a 

13 .2 






°3 
3 

»d 
i 

i> 

a 

bo £ 



1920] 



DUGGAR— H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 39 



3 curves representing the results under the 3 combinations of 
conditions, as well as a fourth curve exhibiting the average of 
the 3 others. 



em 



General Discussion 

Hoagland ('17) determined the effect of H- and OH-ion con- 
centration on the growth of barley seedlings in an incomplete 
nutrient solution, omitting calcium, magnesium, and iron — 

ploying therefore only phosphates of potassium in some 
series and in others adding to these small proportions of sodium 
salts, including nitrate. The H-ion adjustments were made on 
the alkaline side with K3PO4 and K 2 HP0 4 ; on the acid side 
there was used KH 2 P0 4 , supplemented in one case by H3PO4 
and by the dibasic salt. As a result of numerous experiments 
he finds that a concentration of OH-ion greater than 1.8X10 
was injurious, and it was extremely toxic when the concentration 
reached 2.5 XlO -5 . A concentration of H-ion of .7X10 5 was 
favorable to growth while .3 XlO" 3 was very toxic. In these 
solutions there was, of course, opportunity for antagonistic 
effects, and since the solutions were unbalanced, the injurious 
effect of the potassium or sodium ions or both would require 

consideration. 

Some of the complicated effects resulting from the addition of 
salts to toxic acid and alkaline solutions, especially in respect to 
the water relations of plants, have been dealt with by Dach- 
nowski ('14); but inasmuch as the constituents of nutrient solu- 
tions were not involved either in control experiments or other- 
wise the data are scarcely applicable here. 

It is difficult, if not impossible, to attempt a comparison of the 
toxic action of H-ions and OH-ions from the dissocation respec- 
tively of mineral acids and such hydroxides as those of sodium 
and potassium in distilled water with the toxic effects produced 
by the same ions in a culture solution containing diverse other 
ions, especially the cations of the salts usually employed. In 
the latter solutions antagonistic effects, dependent in part upon 
specific relations of the plant employed, must to a certain degree 
obscure the magnitude of the effects. It is of interest to note, 
however, that Kahlenberg and True ('96) found that roots of 
Lupinus albus just lived in n/6400 HC1. Nevertheless, after 5 



40 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



IVol. 7 



days in n/25,600 HC1 the growth rate was but little more than 
one-half that of similar roots in distilled water. Thus we may- 
assume that a hydrogen ion concentration approximately P H 4 
depressed the root growth of this plant. The same seedlings 
survived n/400 potassium hydroxide during 24 hours, but with 
this reagent no growth measurements after an interval of several 
days, comparable to those with HC1, were made. The results 
of Heald ('96) also show that while roots of Pisum sativum 
survived n/6400 HC1, and those of Zea Mays n/1600, still in 
the former the growth rate was low in n/12,800 and for corn 
in n/3200, during an interval of 48 hours. No comparisons 
with distilled water were included. 

Loew ('03), working with the seedlings of Zea Mays, and 
Miyake ('14), using Oryza sativa, have both shown a relatively 
greater resistance of these plants to alkali than to acid, all of 
which emphasizes the importance of devoting special considera- 
tion to the initial acidity of the culture solution. 

Respecting the reaction of soils, there is considerable evidence 
indicating that acidity alone is not necessarily a limiting factor 
in the growth of many crops. With a method considered ade- 
quately accurate as applied to field conditions, Gillespie ('16) 
has examined air-dried samples of 22 crop soils and found the 
P H exponent to vary from 4.55 to 7.1 in the case of 18 soils from 
Maine, Maryland, and Virginia, and a variation of from 8.1 to 
8.7 with 4 soils from Utah and Montana. It is also reported 
by Gillespie and Hurst ('18) that the highest acidity (P„ 4.5) 
was not in the least injurious to potato culture in Caribou and 
Washburn loams, the two main potato-soil types in the region 
in which they worked. 

Plummer ('18), employing the soil-suspension method, ex- 
amined 68 samples of a variety of soil types of humid regions, 
especially of the southern states. Untreated sandy loam or 



exhibited 



3 i.. -, v,i^-6 



hyd 



to 1X10-", while 



i 



Evidence was gained to the effect that the surface film emphasizes 
the direction of the reaction, that is, in acid soils the surface 
film is more acid and in alkaline soils it is more alkaline. This is 



('16). 



Sharp and Hoagland 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 41 



In general, when the reactions of the culture solutions (such 
as have been employed in this work) are strongly acid, the con- 
tact with plant roots effects a change towards neutrality. The 
extent and rapidity of this change, however, depend somewhat 
upon the crop, and especially upon the composition of the cul- 
ture solution. It may be noted, for instance, from tables xi- 
xix that solution A never became neutral, while solution C was 
changed, in the extreme case, to P H 8.6. On the other hand, it is 
not necessarily true that alkaline culture solutions tend to be- 
come acid, as may be seen in the case of solution C. Solution 
B, normally near the neutral point, may be shifted slightly in 
either direction. 

Impelled in part by the general experience of others in field 
work, indicating a general tendency of cultivated soils to become 
acid, Breazeale and Le Clerc ('12) undertook solution-culture 
experiments to determine "the effect of the reaction of the 
culture medium on the growth of wheat seedlings and particu- 
larly on the development of the root," with a view to a possible 
explanation of the results obtained in practical agriculture. 
They regard the acid tendency as due primarily to the decay of 
organic matter and secondarily to the selective action of the 
root; the last mentioned only they proposed to investigate. 
Their experiments were chiefly with certain salts, particularly 
KC1, K 2 S0 4 , and NaNO$, used singly and each in combination 
with solid CaC0 3 . According to their results greater absorption 
of the K ion, when the potassium salts are used, caused the 
solution to become acid; while in the case of the sodium salt, the 
greater absorption of the N0 3 ion tended to produce alkalinity. 
The addition of calcium carbonate precludes the development of 
acid with the potassium salts. In the toxic action reported no 
account is taken of the lack of physiological balance in the 
solutions lacking Ca, and no experiments were made with KN0 3 . 
Moreover, only a titration method (consistent with the irenpral 



alkalinity 



time) was employed in determining acidity and 



Some time previous to this Hartwell and Pember ('07) 



emphasized the " marked prooertv of 
barley, and oats] of renderin 
insufficient, however, to cau 
tions are numerous in the lit 



>f 



Similar observ 



[Vol. 7 



42 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Hoagland and his associates ('17, '18, '19) in a series of articles 
have pointed out some misinterpretations and discrepancies in 
the earlier work, and among other things have shown that with 
certain proportions of salts in solution cultures and in sand cul- 
tures having an initial acid reaction, this reaction was changed 
with the growth of the crops until it was approximately neutral. 
In certain cultures with an initial reaction approximately neutral, 
plants were grown to maturity without change of solution, and 
with the reaction remaining constant throughout. A nutrient 
solution strongly alkaline from the presence of K 3 P0 4 became 
approximately neutral. Hoagland has also emphasized an 
important point appreciated likewise by some earlier investi- 
gators, namely, that the equivalence of positive and negative 
ions in the solutions is maintained, and the state of equilibrium, 
the recognition of which is often too vague, is of necessity kept in 
mind in any discussion of the absorption of ions. 

In this paper it has been pointed out that there is generally a 
decline in growth in solutions A and C when the H-ion concen- 
tration is approximately Ph 6. This may be due to the relative 
insolubility of the phosphates. On the other hand, the generally 
more favorable growth in the B solution at or approaching neu- 
trality may be related in part to the better distribution of phos- 
phate ions or particles, due to the presence of certain substances 
in a state of greater dispersity. In this connection it will be 
recalled that Bonazzi ('19) and Allen ('19) have contributed 
interesting data on the favorable effects of shaking or agitation, 
on the growth of Azotobacter chroococcum, an organism neces- 
sarily grown in alkaline solutions. 

Toole and Tottingham's ('18) results, showing increased 
yield with the addition of ferric hydroxide to Knop's solution, 
are also of particular interest in considering the data obtained 
in this work with solution B. At present there seems to be no 
basis for a final opinion on the r61e of aluminium in promoting 



growth 



The hieh adsorptive property of 



the compound used, together with its buffer action, may be 



erned 



maximum effect of this com 



pound occurred with wheat, but in view of the complication of 
factors involved this may not be significant. 

In the various series of experiments here reported there is 



1920] 

DUGGAR— H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 43 

considerable diversity in the intervals between the renewal of 
solutions. These intervals have varied from four days to a time 
interval covering the entire culture period. The results have 
been as consistent as might be expected, and it is believed that 
the value of lengthening the interval in this way is important 
from the standpoint of reducing the labor in the maintenance of 
such cultures. Trelease and Free ('17) have, however, shown 
that frequent changes of solution are more favorable for growth; 
but it also appears from their data that renewals during the 
first two weeks are not so important as those made later. They 
suggest that a continuous flow of solution through the culture 
is more beneficial than a daily change. 

The work of Pantanelli ('15) and others has shown that after 
plants have been for a few hours in contact with salt solutions 
it may be demonstrated that there has been a different rate of 
absorption of the various ions. The solution, therefore, changes 
rapidly in the presence of abundant absorbing surfaces. Hoag- 
land has also emphasized this point, and within a certain range 
of osmotic concentration he regards the initial concentration of 
any particular ion as practically immaterial. Nevertheless, it 
would be admitted by all that there must be on the one hand a 
true physiological balance, and that on the other hand, the 
concentration of no necessary ion or molecule shall become a 
limiting factor in growth. It is not, however, proposed to dis- 
cuss in this paper the significance and final results of the inter- 
change of ions or molecules between roots and solutions. 

Discussing limiting factors in water cultures Stiles ('16) has 
drawn attention to the limited application of the water-culture 
method in physiological problems. He regards this as related 
to "(1) the difficulty in analyzing results due to the complex of 
factors not under control; (2) the difficulty of controlling in 
some cases even the factor whose action is being investigated; 
and (3) the excess of labor required to produce results which are 
only of a low degree of accuracy." 

In a previous paper Stiles ('15) has in a measure crystallized 
the feeling of many investigators working with water cultures in 
arriving at the conclusion that it is necessary to calculate the 
probable error of the results in accurately evaluating the sig- 
nificance of differences exhibited by different sets of cultures. 



[Vol. 7 



44 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



In a study of probable error he grew single plants in each of 10 
bottles of 1200 cc. capacity, employing uniform methods and 
seed of a selected strain of rye. Four concentrations of nutrient 
solutions were used with the different lots. Experiments con- 
ducted during the early months of the year yielded results as 
follows: The greatest individual variation in any one lot 
amounted to never less than 70 per cent. In the weakest con- 



individual 



H 



the probable error of the mean in these cases was only about 
3-10 per cent of the mean dry weights. Comparable differences 
were found in cultures made later in the spring with a pure line 

of barley. 
That considerable variability has been found by others is 

evident from the examination of the tables in any case in which 

the data have been given in detail. Livingston and Tottingham 

('18) give a table from which it appears that while the culture 

R^a yields only a fair growth of roots the dry weight for the 

entire plant is third and for tops is the highest of all in the 



series. 



however, differs from R 8 C 



one-eighth the concentration of KN0 3 , 3 times as much 
Ca (H 2 P0 4 )«, and 6 times as much MgS0 4 . R 8 Ci is regarded 
as "the best balanced for young wheat plants of all the 
nutrient solutions so far noted in the literature." 

The Shive solution, on the one hand, and the Livingston- 
Tottingham solution, on the other, were of course designed with 
the idea of simplification. The 3 salts employed contain all 
the essential ions except iron. Theoretically, the phosphate, 
nitrate, and sulphate ions may be added in the form of the salts 
of either of the 3 bases or cations, K, Ca, and Mg. However, 
the relatively low solubility of Ca as sulphate may seem to 

render it less practicable to use this salt. There remain 7 
possibilities in the selection of salts, and it might appear that in 
the selection of these the only important points might be, first, 
H-ion concentration, and second, the use of a base required in 
rplativfilv low concentration with an anion which may be sim- 



ply reduced in 
From the em 



known, an 



mi 



consideration is an appropriate ratio between the 
irtion of Ca and Me. Other antagonistic relations 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 45 



also require consideration. An analysis of the differences 
between the Shive and the Livingston-Tottingham solution 
shows that the latter contains a relatively greater concentration 
of K and N0 3 ions and a lower concentration of Ca, Mg, P0 4 , 
and S0 4 . Serious typographical errors in one of their tables 
(table three, page 345) have led Livingston and Tottingham ('18) 
into error in the statement that in the R 5 C 2 solution there are 
2.89 times as many atoms of Ca as of K per unit volume. As a 
matter of fact the partial volume atomic concentration of the 
R 8 Ci solution of Livingston and Tottingham contains more than 
10 times as much K as Ca, while R 6 d contains more than 8 times 
as much K as Ca, and nearly 6 times as much Mg as Ca. The 
Mg : Ca ratio of R 8 Ci is very nearly 2:1. In both of the best 
Livingston-Tottingham solutions the ionic concentrations of 
K and N0 3 are greatest. It seems quite probable that this 
factor, together with the variability in H-ion concentration of 
KH 2 P0 4 , is accountable for the better growth in these solutions. 

Summary and Tentative Conclusions 

The experiments reported in this paper were undertaken 
primarily to determine the influence of variations in hydrogen 
ion concentration on the yield of certain seed plants in solution 
cultures. As the work progressed, however, many modifications 
were suggested, and some of these involved in no way a con- 
sideration of hydrogen ion concentration at points which might 
be regarded as critical for the growth of the crops used. 

The selection of several culture solutions seemed necessary in 
order that some diversity might be introduced in the salt pro- 
portion or composition factors. The solutions employed, and 
their designations, were as follows : solution A, a slight modifica- 
tion of one of Shive's "best" solutions; solution C, a slight 
modification of one of the best Livingston-Tottingham combina- 
tions; and solution B, based in part upon the Crone combina- 
tion of salts, but with this essential difference, namely, that 
"soluble ferric phosphate" was used in place of the "insoluble" 
iron salt. Each of the solutions first mentioned contains a 
monobasic phosphate, and with theoretically pure chemicals 



should 



with a P H exponent about 



Solution B may vary, in my experience, from a hydrogen 



[Vol. 7 



46 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



concentration represented by P H 5.4 to P H 7.1; frequently, 



howev 



,_ 7 

experiments have been carried out in the 



different periods of the year, and represent, on the whole, a con- 
siderable range of environmental conditions. It has been 
impracticable to analyze these except in a very general way, or 
relatively. Wheat and corn have been employed in every series 
of experiments and Canada field peas in all series here reported 

except one. 

Under the most favorable conditions, the 3 solutions men- 
tioned above, without other modification, may all yield excel- 
lent growth. Plants grown in solution B are invariably of a 
deeper green, presenting a finer appearance, and the average 
of the growth quantities (green weight) is higher for wheat and 
corn than in either of the other 2 solutions. In the unmod- 
ified solutions A and C the green weight of peas averages higher 
than in the unmodified solution B. 

Culture solutions prepared with monobasic phosphates may, 
however, exhibit a hydrogen ion concentration which is too high 
for the maintenance of the best growth under certain conditions, 
and especially is this true in the case of wheat. 

Solutions made with monobasic potassium or calcium phos- 
phate free from acid may, under certain conditions, yield max- 
imum growth quantities, but there is often considerable varia- 
bility in the duplicate cultures due to unknown factors. Certain 
grades of the phosphates mentioned— if not specially purified in 
the laboratory— exhibit a P H which may be distinctly toxic. 
Correction of the P H to about 4.8 or 5.2 by means of NaOH or by 
the use, in part, of a dibasic salt generally affords increased 

growth. 

Under extreme conditions— effecting a high evaporation 

rate — it becomes more important to correct to the higher P H 
exponent. Wheat, corn, and peas are sensitive in the order 
named to high hydrogen ion concentration. 

Usually, the addition to solution B of small amounts of dibasic 
potassium phosphate, of solid calcium carbonate, and of alu- 
minium hydroxide has given increased yields, often considerably 
above that of the unmodified solution. The results in the case 
of the aluminium compound are notable in the case of wheat 



1920] 



DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 47 



grown under the conditions described,— related in part, pre- 
sumably, to adsorption and buffer action. 

In general, it would seem that there may be no single "best" 
solution for the growth of any of the 3 plants employed in this 
work. In all probability a ' ' best ' ' solution, like the ' ' optimum ' ' 
temperature, is represented within the "optimum" concen- 
tration rather by considerable range of salt or ion proportions, 
influenced to a greater or less degree by environmental factors. 

If the initial P H of the culture solution is considerably less 
than neutrality there is generally a tendency for this to be 
shifted toward the neutral point, although this depends in part 
upon the composition of the solution and in part upon the plant 
grown. 

I 

Bibliography 

Allen, E. R. (19). Some conditions affecting the growth and activities of Azotobacter 
chroococcum. Ann. Mo. Bot. Gard. 6: 1-44. pi. 1. 1919. 

Bonazzi, A. (19) . On nitrification. II. Intensive nitrite formation in solution. 
Jour. Bact. 4: 43-59. pi. 1. 1919. 

Breazeale, H. F., and Le Clerc, J. A. (12). The growth of wheat seedlings as affected 
by acid or alkaline conditions. U. S. Dept. Agr., Bur. Chem. Bui. 149: 1-18. 
pi 1-8. 1912. 

Burd, J. S. (19). Rate of absorption of soil constituents at successive stages of plant 
growth. Jour. Agr. Res. 18: 51-72. /. 1-13. 1919. 

Clark, W. M., and Lubs, H. W. (17). Colorimetric determination of hydrogen ion 
concentration and its applications in bacteriology. Jour. Bact. 2: 1-34, 109-136, 
191-236. 1917. 

Dachnowski, A. (14). The effects of acid and alkaline solutions upon the water 
relation and the metabolism of plants. Am. Jour. Bot. 1: 412-440. /. 1-4. 
1914. 

Duggar, B. M. (11). Plant physiology. New York, 1911. (See pp. 136-194.) 

, (19). The micro-colorimeter in the indicator method of hydrogen ion 



determination. Ann. Mo. Bot. Gard. 6: 179-181. 1919. 

, and Bonns, W. W. (18). The effect of Bordeaux mixture on the rate of 



transpiration. Ann. Mo. Bot. Gard. 5: 153-176. pi. 10. . 1918. 

, and Dodge, C. W. (19). The use of the colorimeter in the indicator 



method of H-ion determination with biological fluids. Ann. Mo. Bot. Gard. 
6:61-70. f.l. 1919. 

Gillespie, L. J. (16). The reaction of soil and measurements of hydrogen ion con- 
centration. Wash. Acad. Sci. Jour. 6: 71-76. /. 1-2. 1916. 

, and Hurst, L. A. (18). Hydrogen ion concentration — soil type — com- 



mon potato scab. Soil Science 6: 219-236. /. 1-3. 1918. 

Hartwell, B. L., and Pember, F. R. ('07). The relation between the effects of liming 
and of nutrient solutions containing different amounts of acid, upon the growth 
of certain cereals. R. I. Agr. Exp. Sta., Ann. Rept. 20: 358-380. /. 1-2. 1907. 



[Vol. 7 



48 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Heald, F. D. ('96). On the toxic effect of dilute solutions of acids and salts upon 

plants. Bot.Gaz. 22: 125-153. pi. 7. 1896. 
Hoagland, D. R. (17). The effect of hydrogen and hydroxyl ion concentration on 

the growth of barley seedlings. Soil Science 3: 547-560. 1917. 
, (18). The relation of the plant to the reaction of the nutrient solution. 



Science N. S. 48: 422-425. 1918. 

, (19). Relation of the concentration and reaction of the nutrient medium 

to the growth and absorption of the plant. Jour. A gr. Res. 18: 73-117. f. 1-4. 

1919. 
Kahlenberg, L., and True, R. H. ('96). On the toxic action of dissolved salts and 

their electrolytic dissociation. Bot. Gaz. 22: 81-124. 1896. 
Livingston, B. E., and Tottingham, W. E. (18). A new three-salt nutrient solution 

for plant cultures. Am. Jour. Bot. 5 : 337-346. 1918. 
Loew, F. A. ('03). The toxic effect of H- and OH-ions on seedlings of Indian corn. 

Science N . S . 1 8 : 304-308 . 1 903 . 
McCall, A. G. (15). A new metho. 1 for the study of plant nutrients in sand cultures. 

Am. Soc. Agron. Jour. 7: 249-252. 1915. 
(16). Physiological balance of nutrient solutions for plants in sand cul- 



tures. Soil Science 2: 207-254. pi. 1. f. 1-8. 1916. 

-, Norton, J. B. S., and Richards, P. E. (19). Abnormal stem growth of 



soy beans in sand cultures with Shive's three-salt solution. Ibid. 6:479-481. 
P l. 1-2. 1919. 

Miyake, K. (14). Ueber die Wirkung von Sauren, Alkalien und einiger alkali Salze 
auf dem Wachstum der Reispflanzen. Sapporo Nat. Hist. Soc, Trans. 5: 
91-95. 1914. 

Pantanelli, E. (15). Uber Ionenaufnahme. Jahrb. f. wiss. Bot. 56: 689-733. 1915. 

Plummer, J. K. (18). Studies in soil reaction as indicated by the hydrogen elec- 
trode. Jour. Agr. Res. 12 : 19-31. 1918. 

Schmidt, C. L. A., and Hoagland, D. R. (19). Table of Ph, H+, and OH-values 
corresponding to electromotive forces determined in hydrogen electrode meas- 
urements, with a bibliography. Univ. Calif. Publ., Physiol. 5 : 23-69. 1919. 

Schreiner, O., and Skinner, J. J. (10). Ratio of phosphate, nitrate, and potassium 
on absorption and growth. Bot. Gaz. 50: 1-30. /. 1-9. 1910. 

t , (10a). Some effects of a harmful organic soil constituent. 

Ibid. 50: 161-181. /. 1-11. 1910. (See also U. S. Dept. Agr., Bur. Soils Bui. 
70:1-98. pi. 1-4- f.1-31. 1911.) 

, ('11). Organic compounds and fertilizer action. U. S. Dept. 



Agr., Bur. Soils Bui. 77: 1-31. /. 1-5. 1911. 



(12). The toxic'action of organic compounds as modified by 



fertilizer salts. Bot. Gaz. 54: 31-48. f. 1-5. 1912. 

Sharp, L. T., and Hoagland, D. R. (16). Acidity and absorption in soils as measured 
by the hydrogen electrode. Jour. Agr. Res. 7: 123-145. /. 1. 1916. 

Shive, J. W. (15). A study of physiological balance in nutrient media. Physiol. 

Res. i:327-397. f. 1-15. 1915. 
, (15a). A three-salt nutrient solution for plants. Am. Jour. Bot. 4: 



157-160. 1915. 



1920] 

DUGGAR — H-ION CONCENTRATION AND NUTRIENT SOLUTIONS 49 



, ('18). A comparative study of salt requirements for young and for 

mature buckwheat plants in sand cultures. Soil Science 6: 1-32. /. 1-3. 1918. 

, (18a). Toxicity of monobasic phosphate towards soy beans grown in 

soil and solution cultures. Ibid. 5: 87-122. 1918. 



, and Martin, W. H. (18). A comparison of salt requirements for young 

and for mature buckwheat plants in water cultures and sand cultures. Am. 
Jour. Bot. 5: 186-191. 1918. 

Sorensen, S. P. ('09-10). Etudes enzymatiques. II. Sur la mesure et l'importance 
de la concentration des ions hydrog£ne dans les reactions enzymatiques. Lab. 
Carlsberg, Compt. rend, des Trav. 8: 1-168. /. 1-7; 396-401. 1909-1910. 

Stewart, G. R. (18). Effect of season and crop growth in modifying the soil extract. 
Jour. Agr. Res. 12: 311-368. pi H. f. 1-24- 1918. 

Stiles, W. (15). On the relation between the concentration of the nutrient solution 

and the rate of growth of plants in water culture. Ann. Bot. 29: 89-96. 1915, 

? (16). On the interpretation of the results of water culture experiments. 



Ibid. 30: 427-436. 1916. 

Toole, E. EL, and Tottingham, W. E. (18). The influence of certain added solids 
upon the composition and efficiency of Knop's nutrient solution. Am. Jour 
Bot. 5: 452-461. 1918. 

Tottingham, W. E. (14). A quantitative chemical and physiological study of 
nutrient solutions for plant cultures. Physiol. Res. 1 : 133-245. /. 1-15. 1914. 

Trelease, S. F., and Free, E. E. (17). The effect of renewal of culture solutions on 

the growth of young wheat plants in water-cultures. Johns Hopkins Univ. 

Circ. 293:227-228. 1917. 
True, R. H. ('00). The toxic action of a series of acids and of their sodium salts on 

Lupinus albus. Am. Jour. Sci. IV. 9: 183-192. 1900. 
Truog, E. (18). Soil acidity: its relation to the growth of plants. Soil Science 5: 

169-195. 1918. 



4 



HUMIDITY IN RELATION TO MOISTURE IMBIBI- 
TION BY WOOD AND TO SPORE GER- 
MINATION ON WOOD 

SAN FORD M. ZELLER 

Assistant Plant Pathologist, Oregon Agricultural College and Experiment Station; 

Formerly Visiting Fellow in the Henry Shaw School of Botany of 

Washington University 

Introduction 

In a previous publication the writer ('16) pointed out that 
between a certain minimum and maximum of moisture in wood 
Lenzites saepiaria and other similar wood-destroying fungi will 
grow and cause the destruction of the wood. Thus, the power 
of wood to absorb moisture, whether as vapor from the air or as 
water from objects in contact with the wood, is a factor of prime 
importance in its susceptibility to decay. It is no less true that 
any property of the wood which may influence its moisture- 
absorbing capacity is a factor in its durability. In this con- 
nection it has been suggested (Zeller, '16) that, although resin 
has no actual toxic effect on the growth of wood-destroying 
fungi, it does inhibit growth when in large percentages and its 
only inhibitive power probably lies in the fact that it excludes 
water from the fibre containing it (Zeller, '17). 

With these previous results as a foundation further investiga- 
tions have been conducted to ascertain (1) the amount of mois- 
ture which wood will absorb from the atmosphere at different 
relative humidities when the temperature remains constant ; (2) 
whether the water-proofing effect of resin on wood can be meas- 
ured; and (3) the relation of the moisture content of wood (or 
relative humidity of the atmosphere) to the propagation of wood- 
destroying fungi on wood. The purpose of the present paper 
is to report the results of these experiments. 



Experimentation 

Materials used. — For the experiments reported below samples 
were selected from specimens of shortleaf pine (Pinus echinata) 
secured at the Fordyce Lumber Company, Fordyce, Arkansas, and 
from specimens of longleaf pine (Pinus palustris) secured at the 



Ann. Mo. Bot. Gard., Vol. 7, 1920 



(51) 



[Vol. 7 



52 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



sam 



Calciseau Lumber Company, Lake Charles, Louisiana. From 
these samples approximately 250 specimens of each species were 
selected, so that a wide range in resin content and specific 
gravity as well as sap- and heart-wood might be represented. 

The samples were allowed to air-dry for several weeks in a 
room at about 50 per cent relative humidity. They were then 
measured for volume and weighed, and the specific gravity de- 
termined for the whole sample. The samples were usually 
2X6X48 inches. Of course, these specific gravities are not 
standard (Newlin and Wilson, '19), for the specimens were not 
oven-dry when weighed. Since the samples were obtained for 
the purpose of measuring the moisture-absorbing power of the 
wood they were not kiln-dried, for this tends to reduce the 
hygroscopic property of the wood below that obtaining in green 

)les (Tiemann, '07). This point was kindly called to my 
attention by C. H. Teesdale of the Forest Products Laboratory, 
Madison, Wisconsin, and in a report from that laboratory on 
"Wood in aircraft construction" (1919) the relation of kiln- 
drying of wood to its hygroscopicity is quite generally discussed. 
All samples finally selected for the moisture-absorption tests 
were marked and so labeled that after they were cut into blocks 
2X2X4 inches, each block could be identified as to which sample 
it belonged and as to its position in the sample. From these 
small blocks uniformly clear pieces were selected for the prepara- 
tion of shavings which were uniform in thickness. A preliminary 
experiment revealed the fact that the moisture-imbibing power 
of wood is not changed by shaving, but that imbibition is more 
rapid than when blocks were used. Therefore, in the experi- 
ments reported below shavings were used. The clear pieces 
selected were of a light color and had a resin content well below 
5 per cent, except those otherwise designated in tables n and iv. 

Description of humidors. — In order to determine the relation 
between the humidity gradient of the atmosphere and the 
moisture content of wood a closed chamber, or humidor, was 
devised so that a constant temperature would be maintained, 
and so that the humidity could be regulated either by differen t 
concentrations of sulphuric acid in trays enclosed in the chain - 
ber, by varying the evaporating surface of water in trays, or by 
hanging baskets of calcium chloride in the chamber. To pro- 



1920] 

ZELLER — IMBIBITION BY WOOD AND SPORE GERMINATION 53 



duce the higher humidities, curtains of absorbent paper were 
hung in the chamber so that the lower portions were in contact 
with water in trays. This increase in evaporating surface 

proved very effective. 

The humidor with apparatus complete as we used it is shown 
in plate 1. The humidor proper consists of two double-walled 
boxes, one within the other, built of spruce lumber. The 
double walls are packed with sawdust for insulation against 
temperature changes. For added insulation there is an air space 
separating the inner and outer box. The inner box, which is 
the chamber proper, has a double-walled door provided with a 
double-glass window, through which temperature and dew- 
point observations can be made without opening. The outer 
doer is double-walled and packed with sawdust. The doors are 
provided with ordinary cold-storage catches. There are three 
one-inch openings, one through the top and one through each 
end. The inner walls of the humidor chambers were water- 
proofed in two ways: (1) Those of two humidors were brushed 
with hot paraffin (parawax) and were then thoroughly ironed 
with a hot electric iron. Further applications of the paraffin 
were made in the same manner. (2) The walls of two other 
humidors were primed and painted with several coats of a 
water-proof enamel and valspar. The paraffin treatment does 



give as good an a] 
ed to be the better 



enamel paints, but 



temperature of the chamber was maintained at 25° C. by 



means of a bimetallic thermo-reg 
light as a heating element. 
Determination of relative hum 



difficulty 



in maintaining a constant relative humidity throughout any one 
experiment. The relative humidity of the chamber was deter- 
mined by means of a Milliken dew-point apparatus, which 



;hly polished 



ided 



with a three-hole stopper. One of these holes supports a 
thermometer and the other two provide an intake and outlet 
for the siphoning of water or freezing mixtures through the cup. 
This apparatus show T s plainly in fig. 2, pi. 1. The dew-point is 
determined by the appearance and disappearance of the film of 
moisture on the polished cup as the temperature is changed by 



[Vol. 7 



54 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



*• 



allowing water to flow through the siphon. From the vapor 
pressures of the dew-point and the temperature of the chamber 
the relative humidity is obtained. The pressure of aqueous 
vapor at various temperatures was secured by recourse to the 
Smithsonian physical tables (Fowle, '10). 

Method of weighing within the humidor. — The samples of wood 
shavings which were used to measure the imbibition of moisture 
from the atmosphere were planed as needed and placed in the 
wire baskets shown in fig. 1 and 2 of pi. 1. The baskets pro- 
vided with hooks were hung on two wires which were stretched 
across the chamber, one on either side of the openings in the top 
and the right-hand side of the chamber. Balances were in- 
stalled on the top of the humidor in such a position that a wire 
supporting a counterweight passed down through the opening 
leading to the chamber. The lower end of this wire is hooked so 
that the wire baskets can be hung upon the balance by means 
of a lever which passed into the opening in the right-hand end 
of the humidor. This operation can be performed without 
opening the chamber and thus changing the temperature and 
humidity within. The wire lever and the counterbalance wire 
are each provided with a cork, which can be slipped back when 
the device is in use or into the apertures when not in use. This 
device is an adaptation of that described by Dixon ('98). 

A similar one to that described by the writer has been used 

for the same purpose by Mr. C. H. Teesdale in charge of the 

Section of Wood Preservation at the Forest Products Laboratory, 

Madison. 

Determination of moisture content of wood. — After the samples 

of shavings had come to a constant weight in the humidor they 
were weighed and at the same time the relative humidity of the 
chamber was determined and recorded. The samples were then 
dried to constant weight in a dry-air oven at 100-105° C. The 
difference between the weight taken in the chamber and that 
after drying gave the total amount of moisture in the samples 
including that absorbed at the relative humidity of the chamber. 
The per cent of moisture absorbed was based on the oven-dry 
weight of the shavings. 

Determination of resin content of wood. — To determine whether 
the influence of a high resin content on moisture absorption can 



19201 

ZELLER — IMBIBITION BY WOOD AND SPORE GERMINATION 55 



be measured experimentally samples were chosen and analyzed 
by the method previously reported (Zeller, '17). 

Results of Experiments on Moisture Absorption 

The data from the experiments to show the relation between 
the relative humidity of the atmosphere and the moisture con- 
tent both of sap- and heart-wood of shortleaf and longleaf pine 
are given in tables i, n, m, and iv. 

From the data recorded in these four tables (i, n, in, and iv), 
the humidity-moisture curves shown in figs. 1, 2, 3, and 4, 
respectively, were plotted. These curves represent the ulti- 
mate moisture content at a given temperature and relative hu- 
midity of the surrounding atmosphere. The samples used in 
each experiment were of three distinct specific gravities chosen 
to represent the approximate average specific gravity of the 
species of wood, as well as densities both heavier and lighter 
than the average. In the case of heart-wood, however, an extra 
series of samples was used. These were highly resinous and 
consequently very dense. Other than these highly resinous 
pieces, the samples had a resin content well below 5 per cent. 

The four curves are strikingly similar. In all cases where the 
)les of wood represented have specific gravities lying be- 
tween .41 and .75 to .80 and small percentages of resin, the 
general curve for the moisture absorbed is followed up to a cer- 
tain relative humidity where the percentage of moisture taken 
up by the wood of the three densities begins to vary according 
to the density. The lighter, or less dense, samples, from this 
point, take up more moisture with increased atmospheric hu- 
midity than do the denser samples. This occurs at a relative 
humidity averaging from 94.75 to 96 per cent. There seems to 
be but one explanation for a divergence of the curves at this 
juncture. Up to this point the wood fibre has not received 
enough moisture from the atmospheric humidity to satisfy its 
imbibition capacity, but beyond this point this hydration capac- 
ity is over-satisfied and the moisture over and above that ab- 
sorbed by the fibre is adsorbed by the surfaces exposed. If this 
theory is correct the point of divergence of the three moisture- 
humidity curves represents the fibre-saturation point. As an 
irreversible colloid, wood is undoubtedly limited in further 



sam 



56 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



TABLE I 

PERCENTAGE MOISTURE CONTENT OF SHORTLEAF PINE SAP-WOOD SHAVINGS AT 

VARIOUS ATMOSPHERIC HUMIDITIES AND AT 25° C. 



Samples of .41 
specific gravity 


Samples of .61 
specific gravity 


Samples of .69 
specific gravity 


I 


II | III 


IV 


V 


VI 


Moisture 

in shavings 

(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Moisture 

in shavings 

(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Moisture 

in shavings 

(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


4.12 


11.0 


2.5 


6.6 


3.0 


7.5 


5.50 


19.5 


6.62 


24.8 


4.87 


15.25 


7.25 


28.0 7.5 


31.8 


6.12 


22.75 


7.80 


33.2 


7.9 


35.8 


7.88 


34.6 


8.70 


45.5 


8.4 


40.4 


8.70 


42.5 


9.00 


49.5 


9.2 


47.5 


9.7 


53. 


9.88 


53.8 


9.5 


54.5 


10.75 


62.5 


10.62 


58.8 


10.3 


58.8 


11.6 


66.0 


10.80 


60.4 


10 4 


61.3 


12.6 


69.5 


11.12 


63.8 


11.5 


65.0 


14.25 


76.8 


12.80 


68.8 


12 2 


67.5 


15.3 


79.3 


13.88 


73.8 


13 


71.8 


16.3 


84.4 


15.5 


81 


16.2 


82.2 


19.3 


91.0 


17.25 


86.5 


17.6 | 86.4 


23.4 1 95.3 


18.5 


87.8 


20.0 


90.0 


24.5 96.5 


18.75 


90.0 


21.6 


93.2 


25.0 


97.5 


21.4 


92.25 


24.25 




96.0 


26.0 


98.2 


22.3 


94.4 


26.0 


97.4 


26.25 


98.8 


23.8 


95.5 


26 6 i 98.2 


27.1 


99 3 


25.4 

— 


96.6 


27.12 


98 . 2 


27.75 


99.5 


27.6 


97.5 


27.6 


98.7 


28.0 


100.0 


28.9 


98.4 1 28.6 


99.2 






30.25 


98.7 


29.0 


99.5 







31.5 


99.2 


29.88 


100.0 






32.0 99.5 










34.0 100.0 

1 









1920] 



ZELLER — IMBIBITION BY WOOD AND SPORE GERMINATION 



57 



TABLE II 

3 MOISTURE CONTENT OF SHORTLEAF PINE HEART-WOOD 
AT VARIOUS ATMOSPHERIC HUMIDITIES AND AT 25° C. 



Samples of .49 
specific gravity 


Samples of .61 
specific gravity 


Samples of .70 
specific gravity 


Resinous samples of 

.82-90 specific 

gravity 


I 


II J III 


IV 


V 


VI 


VII 


VIII 


IX 


Moisture in 
shaving s 
(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Moisture in 
1 shavings 
1 (per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Moisture in 
shavings 
(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Moisture in 
shavings 
(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


- 

Per cent 
resin 


3.8 


10.3 3.2 


8.0 


3.875 


9.4 


6.62 


24.8 


15.6 


4.75 


13.6 


6.2 


20.4 


5.12 


15.6 


7.9 


33.6 


16.2 


6.5 


23.2 


7.0 


26.2 


6.00 


19.0 


8.5 


38.5 


16.7 


7.2 


30.2 


7.6 


31.8 


7.12 


27.8 


9.32 


43.5 


16.2 


7.5 


33.2 


7.7 


34.6 


7.5 


30.3 


10.2 


49.6 


15.6 


9.0 


41.5 


8.4 


36. 

40.4 


8.4 


37.2 


10.34 


54.4 


18.2 


10.7 


51.8 


8.7 


9.8 


45.6 


11.32 


57.4 


17.6 


12.3 


61.0 


10.1 


44.2 


11.38 


55.8 


12.3 


63.0 


17.6 


14.5 


71.0 


10.2 


47.4 


13.18 


65.8 


13.7 


70.0 


16.7 


16.7 


78.0 


11.2 


53.8 


15.86 


75.6 


13.88 


72.6 


18.2 


18.0 


82.4 


11.8 


59.3 


16.4 


79.2 
80.4 


14.8 


75.6 


15.6 


19.7 


87.5 


13.12 


63.2 
65.0 


17.37 


16.0 


80.4 


18.2 


21.75 


92.0 


12.75 


18.62 


86.0 


16.28 


83.2 


15.6 


23.3 


94.5 1 13.6 


67.5 


20.7 


90.0 17.5 


86.0 


16.2 


24.0 J 95.0 | 15.6 | 72.8 


22.12 


93.0 


18.25 


88.2 


16.2 


25.5 


96.0 16.14 


76.6 


22.75 


94.0 


18.7 


90.6 


15.6 


28.8 


97.8 


17.3 


78.8 


24.25 


95.60 


19.5 


91.4 


18.2 


30.7 


98.4 


18.4 


84.4 


25.2 


97.0 


20.11 


94.4 


17.6 


32.55 


98.8 


19.4 


85.3 28.08 1 


100.0 


20.52 


96.2 18.2 


36.25 


100.0 


19.7 


88.6 






21.4 


96.6 


18.2 






20.3 


S8.8 






21.41 


98.6 


17.6 



[Vol. 7 



58 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



TABLE II— Continued 

PERCENTAGE MOISTURE CONTENT OF SHORTLEAF PINE HEART-WOOD SHAVINGS 

AT VARIOUS ATMOSPHERIC HUMIDITIES AND AT 25° C. 



Samples of .49 
specific gravity 


Samples of .61 
specific gravity 


Samples of .70 
specific gravity 


Resinous samples of 

.82-90 specific 

gravity 


I 


ii 


in 


IV 


V 


VI 


VII 


VIII 


IX 


Moisture in 

shavings 

(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Moisture in 
shavings 
(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Moisture in 
shavings 
(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Moisture in 
shavings 

(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Per cent 
resin 






21.2 


91. 






22.25 


100.0 


17.6 






22.6 


93.2 














... 


24 . 75 


96.2 
















25 75 


97.0 














27.12 


98.0 














29 . 32 


98.8 














31.25 


99.4 
















33.10 


100.0 













1920] 



ZELLER — IMBIBITION BY WOOD AND SPORE GERMINATION 



59 



TABLE III 

PERCENTAGE MOISTURE CONTENT OF LONGLEAF PINE SAP-WOOD SHAVINGS AT 

VARIOUS ATMOSPHERIC HUMIDITIES AND AT 25° C. 



Samples of .41-.42 
specific gravity 


Sample 
specific 


s of .70 
gravity 


Samples of .75-.80 
specific gravity 


i 


II 


III 


IV 


V 


VI 


^ Moisture 

in shavings 

(per cent) 




Relative 

humidity 

of chamber 

(per cent) 


Moisture 

in shavings 

(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Moisture 
in shavings 

(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


3.88 


9.3 J 4.04 


12.3 


5.4 


15.8 


6.78 


28.10 


6.24 


21.4 


6.89 


26.1 


8.24 


37.4 


7.21 


28.2 


7.88 


34.0 


9.52 


47.8 


7.30 


31.4 


8.64 


40.6 


10.83 


56.4 


9.21 


44.7 


9.78 


50.8 


12.2 


64.2 


10.11 


49 4 


10.68 


55 1 


13.77 


71.6 


10.38 


53.6 


11.27 


61.2 


14.82 


75.8 


11 50 


58.6 


12.77 


67.3 


17.59 


83.7 


12.54 


65.8 


15.73 


79.0 


20.31 


88.2 


13.42 


69.3 


19.32 


87.2 


22.38 


92.5 


14.15 


73.8 


21.71 


91.2 


24.14 


94.6 


16.51 


80.3 


23.58 


94.6 


24.88 


95.4 


17.07 


82.5 


23.96 


95.4 


25.76 


96.2 


18.48 


85.5 


24.54 


96.1 


26.63 


96.7 


20.74 


89.7 


' 24.74 


97.2 


28.64 1 97.8 


23.35 


94.2 


25.32 


97.7 


30.52 


98.7 


23.76 


94.6 


26.14 


98.8 


32.18 


99.1 


24.26 


95.3 


27.02 


100.0 


35.76 


100.0 


25.61 


96.8 










26.80 


97.4 










27.63 


98.6 










28 . 59 
30 . 28 


99.1 










100.0 







60 



(Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



TABLE IV 

PERCENTAGE MOISTURE CONTENT OF LONGLEAF PINE HEART-WOOD SHAVINGS 

AT VARIOUS ATMOSPHERIC HUMIDITIES AND AT 25° C. 



Samples of .51 
specific gravity 


Samples of .70 
specific gravity 


Samples of .75- Resinous samples of 
.80 specific gravity | .86-.94 specific gravity 


I | II 


III 


IV 


V 


VI | VII VIII 


IX 


Moisture in 
shavings 
(percent) 


Relative 

humidity 

of chamber 

(per cent) 


Moisture in 
shavings 
(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Moisture in 
shavings 

1 (percent) 


Relative 
humidity 
of chamber 
(per cent) | 


Moisture in 

shavings 

(per cent) 


Relative 

humidity 

of chamber 

(per cent) 


Per cent 
resin 


3.89 


10.6 


3 <is 

5.0 


9.8 
15.0 


4.12 


11.8 

18.6 


6.25 


23.1 
31.2 


19.3 


4.48 


12.6 


5.62 


7.24 


17.1 


5.32 


17.6 


5.88 


20.0 


7.14 


26 . C) 


7.9 


38.3 


18.4 


6.78 


26.0 


7.14 


28.2 


7.16 


29.4 


9.42 


51.6 


16.4 


7.82 


31.6 


7.64 


33.0 


7.8 


34.8 


9.36 


55.4 


19.3 


8.84 


46 . 6 


8 25 


40.0 


8.88 


49.6 


9 71 


58.7 


19.3 


10.08 


54.8 


8.62 


42.2 


9.87 


53.0 


10.58 


58.7 


18.4 


11.06 


59.8 
66.4 


9.28 


48.2 


10.21 


57.6 


10.27 



11.28 


60.8 


17.1 


12.2 


9.31 


50.0 


10.68 


61.2 


64.4 


16.4 


13.8 


74.4 


10.78 


60.0 
63.6 


11.28 


62.5 


11.76 


66.0 


16.4 


15.52 


81.4 


11 64 


11.72 


65.1 


12.26 


68.7 


18.4 


17.10 


85.4 


12.72 


70 


13.38 


72.9 


12.2 


70.8 


16.4 


20.10 


90.8 
93 4 


16 22 


83.8 


14.27 


76.6 


12.81 


71.7 


17.1 


21.78 


17.4 


86.8 


18.54 


88.3 


1 3 02 


74.3 


18.9 


23.68 


95.6 


19.21 


89.4 


19.71 


90 7 


13.94 


78.0 


17.1 


25 1 


96.6 


22.79 


94.6 


20.86 


92.1 


14.31 


78.7 


16.4 


26.62 


97.4 


24 . 36 


96 . 5 1 


23.27 


95 6 


14.82 


82.4 


19.3 


27.8 


98.0 
99 


26.11 


98. 


24.01 


96.5 


15.13 


84.7 


19 3 


30.32 


27 . 57 


99.2 


25 . 24 


97 9 


1 6 . 08 


86 . 9 


17.1 


33.2 


100.0 


29.1 


100.0 


26 11 


99.1 


17.0 


90.3 


19 






26.89 


100 


16.75 

18.48 


91.5 


16.4 














93.5 


19.0 














17.41 


93.8 


19.3 














18.74 95.1 


19.0 














1 9 . 66 


95.3 


10.4 














20 . 1 4 


97.2 


18.4 














20 . 1 5 


99.1 


18.4 














21.12 


100.0 


18.4 



1020] 



ZELLER — IMBIBITION BY WOOD AND SPORE GERMINATION 



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[Vol. 7 



62 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



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1920] 



ZELLER — IMBIBITION BY WOOD AND SPORE GERMINATION 



63 



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MOISTURE 




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ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



PER 



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1920] 



ZELLER — IMBIBITION BY WOOD AND SPORE GERMINATION 65 



absorption of moisture by certain morphological and mechanical 
properties of the woody tissues which vary with the different 
species of wood. This variation in morphological and mechani- 
cal characters in the wood of different species will account for 
the slight differences in moisture content at the fibre-saturation 
point of shortleaf and longleaf specimens. The moisture con- 
tent at the so-called fibre saturation point of the four 'curves is 
approximately as follows: shortleaf pine sap-wood, 24.25 per 
cent, heart-wood, 24.5 per cent, and longleaf pine sap-wood, 
23.75 per cent, and the heart-wood, 23.25 per cent. 

In timber-testing laboratories the fibre-saturation point of 
wood has been determined by means of strength tests, and has 
been defined by Tiemann ('07) as "the degree of moisture at 
which maximum absorption by the cell walls is reached. " After 
this point is reached added moisture does not lessen the strength. 
Beginning with the dry conditions, with the increase of moisture 
the strength falls off very rapidly at first, then more slowly as 
the fibre-saturation point is reached, and here it abruptly ceases 
to decrease. This abrupt break in the moisture-strength rela- 
tion represents the fibre-saturation point. The moisture per 
cent at fibre-saturation for longleaf pine as determined by Tie- 
mann ('07) by compression tests on small specimens averaged 
25 per cent, with a maximum at 26 per cent and a minimum at 
24 per cent. This is not far removed from the moisture content 
at which the curves representing the three different specific 
gravities of longleaf pine diverge in figs. 1 and 2. 

If this divergence of the moisture-humidity curve represents 
the fibre-saturation point the appreciable increase in moisture 
up to 100 per cent humidity must be moisture in the form of a 
surface film; that is, the fibre-saturation point is the point where 
absorption or imbibition by the fibre is replaced by a surface 
phenomenon, adsorption. In a unit weight of wood fibre the 
thin-walled, large-lumened cells, having a lower specific gravity 
than the heavy-walled, small-lumened cells, present a much 
greater surface than the latter. If the greater concave curva- 
ture of the smaller-lumened cells has any tendency to thicken the 
surface film in proportion to that adsorbed by those having less 
curvature, the difference in the total moisture created in this 



dently is not great enough to 
l from the difference in surface 



difference 



5 



[Vol. 7 



66 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



This relation of the moisture content of wood at various rela- 
tive humidities has been demonstrated for other woods by 
workers at the United States Forest Products Laboratory located 
at Madison, Wisconsin. This work is reported in "Wood in 
Aircraft Construction" (1919) which was prepared by the above- 
mentioned laboratory for the United States Navy Department. 
The five woods worked with were Sitka spruce, black walnut, 
white oak, yellow birch, and ash, and the maximum moisture 
content at 100 per cent humidity ranged from about 29.5 to 
30.2 per cent for Sitka spruce to about 36.7 per cent for ash. A 
report by Pfeiffer on the physical properties of several Javanese 
woods includes a study of their moisture contents at various 
atmospheric humidities. The maximum moisture contents of 
the various woods at 100 per cent humidity were as follows: 
Tectona grandis (teak wood), 21 per cent; Ensideroxylon Zwageri, 
22 per cent; Shorea sp., 24 per cent; Dipterocarpus sp., 23 per 
cent ; Shorea sp., 30 per cent, and Alstonia sp., 25 per cent. The 
specific gravities of these woods were .64, 1.04, .88, .64, .41, and 
.34, respectively. Although the order of increase in maximum 
moisture content in these Javanese woods is not in exactly the 
same order as the decrease in their specific gravities, there is a 
tendency in that direction. Of course, a direct relation through- 
out could not be expected because of the great differences in 
morphological and physical structures between various species. 

Resin Content in Relation to Moisture Absorption 

The data on this subject as presented in tables n and iv and 
illustrated graphically in the curves shown in figs. 2 and 4 are 
self-explanatory and for the most part need no discussion. How- 
ever, there are a few points of interest which might be discussed 
briefly. In the lower part of the curves, where the relative 
humidity is less than 50 per cent, the points representing the 
highly resinous samples of wood show no deviation from the 
moisture curves of those samples of wood containing less than 
5 per cent resin. Above 50 per cent humidity, however, 
there is a gradual deviation of the curve representing highly 
resinous specimens. This moisture curve is lower than that 
representing specimens containing less than 5 per cent resin. 
This illustrates the fact that resin actuallv has a water-proof- 



1920] 



ZELLER — IMBIBITION BY WOOD AND SPORE GERMINATION 67 



ing effect upon the wood containing it, and that this water- 
proofing effect is considerable as the fibre-saturation point is 
approached. In the shortleaf heart-wood at a relative humidity 
of 95 per cent the moisture absorption was decreased 15.6 per 
cent by 17.6 per cent resin, and at 100 per cent humidity the 
minimum moisture decrease was 20.8 per cent, and the average 
decrease was 31.5 per cent brought about by 17.6 per cent resin. 
In the longleaf pine heart-wood at a relative humidity of 95 per 
cent a resin content of 16.4-19 per cent had a water-proofing 
effect of 17.4 per cent, while at 100 per cent humidity the water- 
proofing effect of 18.4 per cent resin averaged 29.3 per cent, with 
a minimum of 21.5 per cent reduction in the moisture content. 
Undoubtedly, this water-proofing effect of resin has its in- 
fluence upon the durability of structural timber placed under 
very humid conditions, providing the resin content is sufficient 
to lower the moisture content of the wood below that which is 
conducive to the growth of wood-decaying fungi. Although the 
resin does have some influence in this direction, it is probably not 
sufficient to be relied upon as a test of durability. More reliance 
could be put upon resin as an index of durability in timbers con- 
taining, for example, 12 to 15 per cent resin, providing there was 
any reason to believe that it was equally distributed throughout 
the timber. With such an equal distribution of the resin the 
water-proofing would undoubtedly be more effective at lower 
relative humidities. Not only this, but mechanical resistance 
of resin equally distributed would undoubtedly be a great factor 
in the inhibition of fungous growth in the wood. This, however, 
is not the case. The resin is deposited in streaks in wood so 
that there are portions relatively free from resin. These 
resin-free portions, having a hygroscopicity sufficient to take 
up considerable moisture under the right moisture conditions, 
become sources of weakness because of the inroads of wood- 
destroying fungi. Examples of such decay are often reported 
as very destructive to the ceilings and structural timbers under 
the highly humid conditions produced in the paper- and pulp- 
mills of the eastern states and Canada and the knitting-sheds of 
the cotton industries of New England. Further experimenta- 
tion along this line is advisable. Some practical process of 
treating resinous lumber, possibly by modifying the kiln-drying 



[Vol. 7 



68 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



processes so as to distribute the resin more evenly, is a worthy- 
project of investigation of this important problem in these 



indust 



Spore Germination on Wood 



The spores of wood-destroying fungi exhibit a marked simi- 
larity with regard to their requirements for germination. Some 
of the factors which influence germination are proper tempera- 
ture and proper amounts of acidity and moisture. As early as 
1860 Hoffmann ('60) found that in a period of five days spores 
of Polyporus versicolor germinated in moist air, and in a period 
of six days spores of P. squamosus germinated better in moist air 
than in water. On the other hand, Falck ('09) says that the 
spores of Lenzites on wood germinate only when the wood has 
been thoroughly saturated by rains. 

To determine more accurately the relation of moisture to the 
germination of spores of a wood-destroying fungus the following 
experiments were conducted, using the spores of Lenzites saepia- 
ria. The spores of this fungus were obtained from rejuvenated 
fruiting bodies as described in a previous paper (Zeller, '16). 
The spores were allowed to drop from the fruiting bodies directly 
upon thin shavings of shortleaf pine (Pinus echinata) sap-wood, 
or were caught in Petri dishes and transferred to the shavings in 



distilled water and allowed to dry rapidly 



air. 



The shavings upon which the spores were deposited were 
clamped in a small device which may be described as follows: 
Two pieces of sheet celluloid, about the size of a microscope 
slide, were riveted together at one end by means of a paper rivet, 
and a round hole about 1 cm. in diameter was cut through the 
middle of the two celluloid slides. The pieces of celluloid were 
bent apart while a shaving to which the spores were adhering 
was slipped between, the other end of the pieces of celluloid 
being clamped together by means of a paper clip. The shaving 
is thus held so that it covers the hole in the middle of the device 

1 A decoction from 60 gms. of shavings of shortleaf-pine sap-wood was prepared b y 
steaming the shavings in a reflux with 10 cc. of distilled water. This decoction teste d 
P H 4.2, so that the actual active acidity of the shavings probably was not above 
P H 3.8. This would be well toward the optimum acidity (P H 3.1) for the germination 
of the spores of Lenzites saepiaria as reported by Webb ('19). 



1920] 



ZELLER — IMBIBITION BY WOOD AND SPORE GERMINATION 



69 



and is exposed to the air from both sides. These devices were 
numbered, and after the chamber of the humidor, described 
elsewhere in this paper, had come to a constant humidity, one 
of them was lowered into the chamber through the opening in 
the top. By means of the lever it was hung on a wire without 
opening the chamber. In this way spores were exposed to the 
atmospheric conditions of the chamber without coming in 
contact with any other object. A new shaving was added 
to the chamber on 12 successive days. All were removed at 
one time, examined under the microscope, and the percentage 
of germination estimated on the basis of the total number of 
spores in a unit field. In table v the percentages of germination 



TABLE V 

RELATION OF THE GERMINATION OF THE SPORES OF LENZITES SAEPIARIA TO 

THE RELATIVE HUMIDITY OF THE ATMOSPHERE AT 25° C. 



Germination 
period (days) 


Per cent 
germination 

(averages) 


Per cent 

relative 

humidity 


Germination 
period (days) 


Per cent 
germination 

(averages) 


Per cent 

relative 
humidity 


10 


1.5 


63.0 


5 


76.0 


96.5 


10 


2.5 


65.0 


5 


85.5 


98.5 


10 


3.5 


67.4 


5 100.0 


99.0 


10 1 2.5 * 


72.5 


5 


100.0 


98.0 


7 4.0 


78.5 


5 


95.0 


98.0 


7 


7.5 


82.5 


6 


57.5 


96.0 


7 


5.5 


85.5 


6 


80.0 1 98.0 


7 


10.0 


89.8 


6 


87.5 


98.0 


5 20.5 


90.5 


6 


92.0 


99.0 


5 


25.5 


92.5 


6 


96.5 


99.0 


5 70.0 


95.5 


6 


100.0 


98.5 



are the averages of three or four counts. In fig. 5 the relation of 
spore germination to relative humidity is presented graphically. 
This curve indicates that until the fibre-saturation point is 
reached the percentage of spore germination is very low, but as 
soon as the humidity of the air is sufficiently high to supply free 



70 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 




60 



70 



80 



90 



100 



PER 



CENT HUMIDITY 



Fig, 5. Curve showing the percentage of germination of spores of Lenzites saepiaria 
on shavings of the sap-wood of Pinus echinata at various atmospheric humidities 
and at 25° C. 



1920] _, 

ZELLER — IMBIBITION BY WOOD AND SPORE GERMINATION 71 



water as a film on the wood surface the spores of Lenzites sae- 
piaria show a high percentage of germination. 

By growing Ceratostomella coerulea upon blocks of pine sap- 
wood Munch ('09) found that free-water on the wood was 
necessary to sustain growth. Growth is maintained if fibre- 
saturation is maintained, but if the water of imbibition falls 
below the saturation point the wood seems to demand water at 
the expense of the mycelium. Although true for Ceratostomella 
this very probably is not true for the mycelium of such fungi as 
Lenzites saepiaria, Merulius lacrymans, or Coniophora cerebella, 
which after becoming well established, seem to maintain a 
water supply from some unknown source. This is especially 
the case with Merulius lacrymans. Wehmer ('14) found that 
this fungus would not grow on blocks of wood in ordinary cellar 
humidity unless the blocks were first saturated with water. 
After the growth was well established, however, the water con- 
tent of the wood was maintained by the fungus above that in 
sound wood under the same conditions. 

It would seem then that for the germination of the spores and 
the establishment of the mycelium of wood-destroying fungi, 
the wood must contain enough moisture to saturate the wood 
fibre. This becomes a serious problem then in such buildings as 
paper- and pulp-mills and knitting factories where high humidi- 
ties are maintained. If the temperatures fluctuate across the 
dew-point, the moisture content of any exposed woodwork is 
maintained up to fibre-saturation and the wood is sure to decay 
unless some method of treatment of the timbers is possible. 



Summary 

In this paper are reported the results of experiments (1) 
showing the moisture content of wood at various atmospheric 
humidities and at 25° C. Curves are presented to illustrate this 
relation for the sap- and heart-wood of both longleaf and short- 
leaf pine. 

(2) By testing the moisture content of any one species of wood 

samples having various specific gravities at the various humidi- 
ties it is possible to approximate the fibre-saturation point of 

the wood. 

(3) The moisture-humidity curves of highly resinous samples 



_. [Vol. 7 

72 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



of wood illustrate the water-proofing effect of resin on the wood, 
especially above 50 per cent humidity. It is believed that 
resin cannot be relied upon as an indication of the durability of 
lumber although present in amounts as high as 12-15 per cent, 
for it is seldom equally distributed in the wood. The regions of 
low resin content are sources of weakness if wood-destroying 
fungi gain access to them under favorable moisture conditions. 

(4) The germination of the spores of Lenzites saepiaria on 
wood shavings was accomplished at various relative humidities 
ranging from 63 to 100 per cent. The germination curve illus- 
trates the fact that spore germination is exceedingly accelerated 
when the atmospheric humidity is high enough to maintain fibre 
saturation of the wood. 

(5) A humidor for maintaining constant humidity and temper- 
ature is described. It is provided with (1) a dew-point appa- 
ratus for the determination of humidity, and (2) a weighing 
device so that the samples can be weighed without opening the 
humidity chamber. 

The writer wishes to express his appreciation of the financial 
aid accorded him by the Southern Pine Association, without 
which this work would have been impossible. Thanks are also 
due the staff of the Missouri Botanical Garden for cooperation 
and for facilities for the work, and to Dr. Hermann von Schrenk 
for helpful suggestions. 



Literature Cited 

Dixon, H. H. ('98). On the effects of stimulative and anaesthetic gases on transpira- 
tion. Trinity College Botanical School, Notes 1898: 97-105. /. 1. 1898. 

Falck, R. ('09). Die Lenzites-faule des Coniferholzes. Moller's Hausschwammfor- 
schungen. Heft 3: 1-234. pi 1-7. f. 24, Jena, 1909. 

Fowle, F. E. ('10). Smithsonian physical tables. Smithsonian Misc. Coll. 58 1 : 1- 
318. 1910. (See tables 135 (d) and 136.) 

Hoffmann, H. ('60). Untersuchungen uber die Keimung der Pilzsporen. Jahrb. f. 
wiss. Bot. 2 : 267-337. pi. 26-35. 1860. 

Munch, E. ('09). Untersuchungen uber Immunitat und Krankheitsempfanglicht- 
keit der Holzpflanzen. Naturwiss. Zeitschr. f. Forst u. Landw. 7: 54-75, 87- 
114, 129-160. /. 1-5. 1909. 

* 

Newlin, J. A., and Wilson, T. R. C. ('19). The relation of the shrinkage and strength 
properties of wood to its specific gravity. U. S. Dept. Agr. Bui. 676: 1-35. 
/. 1-9. 1919. 



1920] 



ZELLER — IMBIBITION BY WOOD AND SPORE GERMINATION 73 



Pfeiffer, J. P. ('17). De Waarde van Wetenschappelijk Onderzoek voor de Vaststel- 
ling van Technische Eigenschappen van Hout. pp. 1-289. pi. 1-4. f. 1-38. 
Amsterdam, 1917. 

Tiemann, H. D. ('07) . The strength of wood as influenced by moisture. U. S. 
Dept. Agr., For. Serv. Circ. 108: 1-42. /. 1-6. 1907. 

Webb, R. W. ('19). Studies in the physiology of the fungi. X. Germination of the 
spores of certain fungi in relation to hydrogen ion concentration. Ann. Mo. 
Bot. Gard. 6: 201-222. /. 1-5. 1919. 

Wehmer, C. ('14). Hausschwammstudien IV. Versuche ueber die Bedingungen 
der Holzansteckung und -Zersetzung durch Merulius. Myc. Centralbl. 3: 
321-332. /. l.;Ibid. V.4: 241-252. f.l; 287-299. pi 1-2. 1914. 

Zeller, S. M. ('16). Studies in the physiology of the fungi. II. Lenzites saepiaria 



Fries, with special re 
512. pi. 8-9. 1916. 



Ann. Mo. Bot. Gard. 3: 439 



, C17). Ibid. III. Physical properties of wood in relation to decay 

induced by Lenzites saepiaria Fries. Ibid. 4: 93-164. pi. 9-13. /. 1. Charts 
1-11. 1917. 



[Vol. 7, 1920] 



74 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Description of Plate 



PLATE 1 

Fig. 1. A general view of the humidor as described on page 53, with the balances 
in place and both doors open. 

Fig. 2. The interior of the humidor showing the baskets containing the samples 
of shavings, one basket hanging on the balance and the Milliken dew-point apparatus 
in the central foreground. 



Ann. Mo. Bot. Gard., Vol. 7, 1920 



Plate 1 




^ 













Fig. 1 




Fig. 2 



ZELLER, IMBIBITION BY WOOD, AND SPORE GERMINATION 



ON CARBOHYDRATE CONSUMPTION BY 

AZOTOBACTER CHROOCOCCUM 

E. R. ALLEN 

Formerly Associate in Biochemistry, Washington University School of Medicine 

and Visiting Investigator, Missouri Botanical Garden 

In a previous paper (Allen, '19) devoted mainly to the study 
of the cause of beneficial action resulting from mechanical agi- 
tation of Azotobacter solution cultures, the defects of existing 
experimental methods for the study of the physiology of this 
organism were pointed out. The present report covers a con- 
tinuance of these studies aimed at the following improvements : 
(1) renewal of the energy source in the cultures; (2) simultaneous 
determinations of nitrogen and of residual carbohydrate at short 
intervals; and (3) mechanical improvements. 

The object of the first of these improvements was, of course, 
to produce heavier growths in the cultures and therefore more 
marked changes in the amounts of metabolic products, thus 
increasing the reliability of the measurements of such changes. 
In view of the theoretical considerations of Duclaux ('98-'00) in 
regard to the rate of increase in the number of cells, we were jus- 
tified in expecting a mounting rate of total physiological activity 
as growth proceeded, until the normal rate of development was 
checked by the accumulation of unfavorable by-products. 
The method of renewal of energy-supplying material has been 
used, for example, with marked success by Bonazzi ('19) in his 
investigations of the organisms of nitrification. 

The second improvement was designed to furnish a more com- 
plete picture of growth processes of Azotobacter than is obtainable 
by the determination of one, or even two, metabolic products at 
the end of an arbitrarily chosen incubation period. Unfortu- 
nately, the "micro" determination of carbohydrate and nitrogen 
on the same sample proved more difficult than was expected, and 
had to be postponed for the time being. The present work, 
therefore, lacks this improvement over that reported previously. 

The third improvement was to facilitate experimental manip- 
ulation and possibly obtain even better results from mechan- 
ical agitation. The type of agitation decided upon was a slow 



Ann. Mo. Bot. Gard., Vol. 7, 1920 



(75) 



(Vol. 7 
76 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



rate of revolution of round-bottomed culture flasks held in a hori- 
zontal position. A rotating device was constructed, which 
held a vertical wooden wheel, to the rim of which were attached, 
horizontally, four one-liter round-bottomed flasks. The wheel 
revolving at the rate of one revolution in five minutes kept the 
culture medium in a slow, uniform motion. 

Only one experiment was conducted, and, as pointed out 
above, it was lacking in the second improvement. Carbo- 
hydrate only was determined at short intervals, and the only 
renewal of this material was in two of the flasks on the rotator. 
The results obtained were so consistently contrary to expectation 
that the data from even the one experiment is of some interest; 
that is, as measured by carbohydrate consumption, the cultures 
showed a declining instead of an increasing metabolic activity. 

The Azotobacter culture used was a later generation of the 
same strain used in previous work. The nutrient solution was 
prepared as follows: 

Solution A: 



Dipotassium phosphate 

Sodium chloride 

Dextrose 



0.2 gm. 

0.2 gm. 

20.0 gms. 

Water, distilled 500 .0 cc. 



Solution B: 



0.2 gm. 
0.1 gm. 



Magnesium sulphate 

Calcium sulphate 

Water 500 .0 cc. 

Ferric chloride (10 per cent sol.) 3 drops 

Solutions A and B were mixed, and 100-cc. portions then 

placed in each of four one-liter round-bottomed flasks and in 
each of two two-liter Erlenmeyers, all flasks receiving in addi- 
tion 1 gm. calcium carbonate. The flasks were then plugged 
with cotton and capped with beakers in the usual way. After 
sterilization by the intermittent method, all except one round- 
bottomed flask were inoculated with a spiral of a suspension 
prepared from an agar slant, the round-bottomed flasks placed 
in position on the rotator, and the Erlenmeyers on a shelf near 
by. The whole experiment was set up in a warm room kept at 
28-30° C. by a thermostat. 

The experiment was started March 14, 1919. Two days later 
a distinct turbidity appeared in the inoculated flasks on the 



1920] 



ALLEN — CARBOHYDRATE CONSUMPTION BY AZOTOBACTER 



77 



rotator, and a lesser one in the Erlenmeyers on the shelf. At 
short intervals 5-cc. samples were removed aseptically and sub- 
mitted to duplicate sugar determinations by the modified 
Shaffer method described in the previous paper (Allen, '19). 
Cultures II and IV received on March 24, 25 cc. of an 8 per 
cent dextrose solution. Culture No. Ill was unfortunately 

lost at this point. 

The data are reported as dextrose remaining per 100 cc. of 
culture solution, no correction being made for evaporation; that 



if 



5-cc. sam 



is found to contain 35 mgs., the 



reported is 0.7000 gm., regardless of the total volume of the 

culture at the time. 

The results appear in tabular form in tables i and n, and in 

graphical form in fig. 1. 



TABLE I 

RESIDUAL DEXTROSE PER 100 C 

DIFFERENT PERIODS 



Culture no. 



March 17 



March 19 



March 24 



March 28 



April 2 



Rotator: (round-bottomed flaaks) 



I (Check) 



II 
III 

IV 



2 . 1565 
1 . 2732 
1 . 4822 
0.9410 



. 7600 
1 . 3507 
. 5700 



0.110-2.060 
. 908-Lost 
. 069-2 . 023 



1.976 



1.842 



1.255 



1.358 



Shelf: (Erlenmeyer flasks) 






V 
VI 



1 . 7257 
1 . 5962 



1.636 
1.418 



1.674 
1.507 



1.523 
1.389 



TABLE II 

MILLIGRAMS DEXTROSE CONSUMED PER DAY IN EACH PERIOD 



Culture 


First 


Next 


Next 


Additional 


Additional 


no. 


3 days 


2 days 


5 days 


4 days 


5 days 


I Check 












II 


295 


256 


130 


21 


144 


III 


225 


65 


88 






IV 


405 


185 


100 


45 


97 



78 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



In constructing the graphs the curves were plotted in the 
normal manner, with the amounts of dextrose on the Y axis 
and time on the X axis for the first ten days of the incubation 
period. In order to make the curves for cultures II and IV 
continuous and harmonic, the Y ordinate is shifted for these 




days 

Fig. 1 



cultures at the end of 10 days so that the 2000-mgm. point is 

on the original X axis. The dotted curve in the insert is the 

Duclaux curve for increase in number of cells, while the solid 

line represents the type of curve predicted on the assumption 

that carbohydrate consumption is a simple function of cell 
multiplication. 

The results show that the rate of carbohydrate consumption 
in Azotobacter cultures does not proceed in a manner similar to 



1920] - ft 

ALLEN— CARBOHYDRATE CONSUMPTION BY AZOTOBACTER 79 

the rate of increase in cell numbers predicted by Duclaux for 
bacterial cultures in general. Indeed, the curves resemble the 
antipode of the Duclaux curve. 

It might be argued that the decrease in rate of sugar consump- 
tion with increase of time in the cultures is due to accumulation 
of by-products, and that this decreased activity corresponds to 
the falling away from the purely mathematical curve in older 
cultures, as discussed by Duclaux. The production of toxic 
substances within seven days in cultures as slow growing and as 
dilute as these seems to us to be unlikely. Moreover, the fact 
that the cultures are able to utilize a second portion of sugar 
seems to us to be opposed to the idea of growth-inhibiting sub- 
stances in the medium. 

The results show also that the cultivation of the organism 
under the influence of mechanical agitation of a certain type 
influences to a marked extent the carbon assimilation in its time 

relations. 

On the whole, the experiment emphasizes the need for more 

studies on the periodic changes in the culture of this organism. 



In conclusion, the writer wishes to express his thanks to Pro- 
fessor P. A. Shaffer for rendering this work possible; to Professor 
B. M. Duggar for suggesting the method of attack, and to Mr. 
A. Bonazzi for suggestions in regard to the arrangement of the 
manuscript. 

Literature 

Allen, E. R. ('19). Some conditions affecting the growth and activities of Azoto- 
bacter chroococcum. Ann. Mo. Bot. Gard. 6: 1-44. pi. 1. f. 1-2. 1919. 

Bonazzi, A. ('19). On nitrification. II. Intensive nitrification. Jour. Bact. 4: 
43-59. pl.l. f.l. 1919. 

Duclaux, E. ('98-'00). Traite de microbiologic. 1: 208-212. 1898; Ibid. 2: 
60-75. 1900. 



Annals 






of the 



Missouri Botanical Garden 



Vol. 7 APRIL-SEPTEMBER, 1920 No. 2-3 



THE THELEPHORACEAE OF NORTH AMERICA. XII 1 

• Stereum 



EDWARD ANGUS BURT 

Mycologist and Librarian to the Missouri Botanical Garden 

Professor in the Henry Shaw School of Botany of 

Washington University 

STEREUM 

Stereum Persoon, Roemer Neues Mag. Bot. i:110. 1794; 
Obs. Myc. i:35. 1797, and 2:90. 1799; Fries, Obs. Myc. 
i:274. 1815, Gen. Hym. 14. 1836, Epicr. 545. 1838; 
Hym. Eur. 638. 1874; Berkeley, Brit. Fung. 270. 1860; 
Morgan, Cincinnati Soc. Nat. Hist. Jour, io: 193. 1888; 
Sacc. Syll. Fung. 6:551. 1888; Massee, Linn. Soc. Bot. Jour. 
27:158. 1890; Engl. & Prantl, Nat. Pflanzenfam. (i:l**): 
123. 1898.— B. Sterea of Thelephora, Schweinitz, Naturforsch. 
Ges. Leipzig Schrift. 1 : 105. 1822.— ****Stereum of Thele- 
phora, Persoon, Myc. Eur. 1 : 116. 1822. — Includes Podoscypha 
Patouillard in Duss, Fl. Crypt. Antilles Fr. 230. 1904. 
Includes Lloydella Bresadola in Lloyd, Myc. Writ, i . Myc. 
Notes 6:51. 1901; Sacc. Syll. Fung. i6:1116. 1902. 
Includes Bresadolina Brinkmann, Ann. Myc. 7: 289. 1909. 

Fructifications coriaceous to hard, stipitate, dimidiate or 
effuso-reflexed ; hymenium inferior, not containing setae; inter- 
mediate layer of longitudinally arranged hyphae normally 
present; basidia simple; spores white, even — rough in but few 
instances. 

The species mentioned or described as belonging in Stereum 

1 Issued Dec. 8, 1920. 



Ann. Mo. Bot. Gabd., Vol. 7, 1920 



(81) 



[Vol. 7 

82 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



upon its publication are Stereum hirsutum, S. striatum, S. 
purpureum, S. nitidum, and S. rugosum, no one of which was 

designated as the type species. 

The species of Stereum are here arranged in the usual sections 
of central-stemmed, lateral-stemmed, merismatoid, and dimid- 
iate and effuso-reflexed species; these sections are convenient 
for locating species approximately, but one should bear in mind 
that some species are ambiguous with regard to sectional charac- 
ters; all the species are probably so variable that individuals 
may be selected from most gatherings which will prove very 
misleading for study. For example, Stereum fasciatum is 
properly included in the section of effuso-reflexed species, yet 
fructifications of this species do occur now and then with elonga- 
tion of the umbo so great as to lead one to regard such a fructifi- 
cation as lateral-stemmed. 

While Stereum is a large genus in the number of its North 
American species, its difficulty is not proportional to the number 
of species, for the species of each of its several sections differ 
among themselves microscopically in the absence or presence 
of definite recognizable organs or combinations of organs, such 
as conducting organs containing latex (milk), vesicular organs, 
gloeocystidia, cystidia of various kinds, and noteworthy para- 
physes. In the determination of any species, one's effort is 
soon concentrated upon a small group of four or five species 
of common structure, some of which may be eliminated by 
geographic range, spore dimensions, etc. The structural fea- 
tures have been very important in working out the extensive 
multiplication of species which had arisen in this genus through 
disregard of the work of earlier mycologists. 

As heretofore noted in the case of Hymenochaete, the east and 
west range of the species of Stereum is marked in comparison 
with north and south range ; of our 77 species, only 7 range over 
both north temperate and tropical areas; the other 70 may be 
arranged in two groups, of which the 29 species comprising the 
northern group are in the region from Canada to the Gulf states; 
the other 41 species range from the Gulf states southward. The 
Gulf states are a region in which northern and southern species 
overlap in range. The excess of tropical and subtropical species 
over northern species is due to the small number of northern 



19201 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 83 



stipitate and merismatoid species, of which we have only 5 as 
against 23 in the warmer southern region. The stipitate and 
merismatoid species grow sometimes on dead wood and some- 
times on the ground; all 49 dimidiate and effuso-reflexed species 
grow on dead wood, causing its decay, and are distributed 24 
in the northern and 18 in the southern area, while 7 others are 
the species already mentioned as ranging over both north 
temperate and tropical areas. 



Key to the Species 

§1. Central-stemmed species. — Pileus more or less infundibuliform, some- 
times deeply split on one side, usually stipitate; stem typically central 
or eccentric but lateral-stemmed forms are also present in many of the 
species 1 

§11. Lateral-stemmed species.— Pileus dimidiate, flabelliform, or wedge-shaped 

-—never infundibuliform — attenuated at the base into a more or less 

distinct stem 9 

§111. Merismatoid species. — Pileoli several, somewhat infundibuliform, wedge- 
shaped, or strap-shaped, borne on or along a common stem 12 

§IV. Sessile species, wholly lacking stem or stem-like base.— Pileus dimidiate- 
sessile, umbonate-sessile, or reflexed, all growing on wood — many 
typically reflexed species may sometimes occur wholly resupinate 13 

§1. CENTRAL-STEMMED SPECIES 

1. Fructifications solitary or gregarious 2 

1. Fructifications cespitose ...... 8 

2. Species with pileus always more or less infundibuliform, lacking dimi- 
diate or other lateral-stemmed forms 3 

2, Species having lateral-stemmed forms occurring more or less frequently 

in collections 5 

3. Neither cystidia nor gloeocystidia present; stem not radicated 4 

3. Gloeocystidia present; growing on the ground, l£-3 cm. high, 3 mm.-2 cm. in 

diameter; in South Carolina to Brazil 3. S. Ravenelii 

3. Gloeocystidia present; growing on wood; in West Indies to Dutch 

vjUl£LIl£i ... # # • • « A m $• SU7*Z7lQ,77l€TlS€ 

3. Hair-like cystidia present; pileus white, 2-4 cm. high; in New York to Missouri, 

and in Alabama, Washington and California 10. S. diaphanum 

3. Hair-like cystidia present; pileus slightly darker than S. diaphanum, 3-5 

mm. high; in New York u. S. exiguum 

3. Hymenial organs unknown; growing on the ground, with stem continued 

by a long radicated portion which penetrates deeply; in French Guiana 

* 5. S. macrorrhiza 

4. Growing on wood, 2-15 cm. high and in diameter; upper surface with 

raised, radial ridges; in Gulf states to Bolivia LS. caperatum 

4. Growing on wood, 6-11 cm. high and in diameter; upper surface not 

ridged; pileus and stem'velvety ; in South America. . .#. S. hydrophorum 

5. Neither cystidia nor gloeocystidia present; pileus cartridge-buff to pinard- 
yellow when fresh; in New Hampshire to North Carolina and Tennessee, 
and in Japan 6. S. Burtianum 

5. Hymenial organs unknown; pileus "straw-colored," 1J mm. in diameter; 
stem 4 mm. high; growing on wet ground among moss in Cuba 

m _. • • • • 7. S. rivulorum 

5. Cystidia present g 

5. Gloeocystidia present; no cystidia 7 

6. Pileus white, of soft, bibulous texture, 3-5 mm. broad, 5-7 mm. long; 

on bark and mosses in Cuba 8. S. quitquiliare 



[Vol. 7 



84 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



6. Pileus drying bright yellow, finally fading in the herbarium, of bibulous 

texture; in West Indies to Paraguay ; .9. S. aurantiacum 

6. Pileus drying tawny olive to Saccardo's umber, not of bibulous texture 
but coriaceous-hard instead; lateral-stemmed forms are the more 
common; 5 mm -2$ cm. high, 2-10 mm. broad; in New York to 

Cuba, and in Wisconsin 12. S. tenerrimum 

7. Somewhat cespitose, obscurely zonate, not bearing a cluster of coarse 

processes near base of the pileus, 1^-4 cm. high, 8 mm.-3 cm. in 
diameter; in Ohio and North Carolina to Mexico and West Indies 
IS. S. pergamenum 

7. With a crest of coarse hairs or processes towards base of the pileus; pileus 

6-10 mm. across; on dead Vitis in South Carolina 14. S. cristatum 

8. Hair-like cystidia more or less numerous but no gloeocystidia ; pileus 
cartridge-buff, strigose-squamose; on the ground, Vermont to 

North Carolina, and in Europe 15. S. pallidum 

8. Gloeocystidia barely distinguishable, but no cystidia; pileus plicate on 
both surfaces, with the upper diamine-brown and the hymenium 
white; on the ground, Porto Rico to British Guiana. . .16. S. elegans 
8. Gloeocystidia present but no cystidia; pileus drying pale cinnamon; 

on dead wood, Jamaica to Trinidad 17. S. decolorans 

§11. LATERAL-STEMMED SPECIES 

9. Fructifications not cespitose 10 

9. Fructifications cespitose; pileate segments pectinate along their margins; 

on decaying wood, Carolina to Bolivia .24. S. Hartmanni 

9. Fructifications rarely cespitose, usually gregarious; margin of pileus thick 

and entire; spores 6X5/x, becoming subangular; in Jamaica to Dutch 

Guiana 18. S. radicans 

10. Growing on ground; pileus white when fresh, drying smoke-gray, 

not zonate; spores 4-5^ X 3-5m! * n West Indies 19. S. pusiolum 

10. Growing on dead wood; pileus of soft bibulous texture, drying pinkish 

buff, 3-6 mm. wide, 5-7 mm. long; in Cuba and Porto Rico 

23 S. cyphelloides 

10. Growing on dead wood; pileus not of soft bibulous texture 11 

11. Pileus drying Verona-brown to chestnut, minutely velvety; stem velvety; 

spores 4-5X3-4 M ; in the West Indies 20. S. glabrescens 

11. Pileus whitish when living, livid and pellucid upon drying, 4-6 cm. high, 

with stem ^-1 cm. long; in Guadeloupe 21. S. flabellatum 

11. Pileus white when fresh, drying " reddish brown," 8-15 mm. long, 3-15 mm. 

broad, often deeply split into segments; in Brazil 22. S. fissum 

§111. MERISMATOID SPECIES 

12. Densely cespitose and concrescent throughout into a cluster 7 cm. in 

diameter, with color and aspect of Tremellodendron pallidum; in 
Mexico and Dutch Guiana . . .25. S. craspedium 

12. Fructification a sessile, rosette-shaped mass of reddish brown pileate 

flaps; in San Domingo . ... .26. S. petalodes 

12. Fructification stipitate, white, with many pileate divisions extending 

out from a common stem ; with aspect of doubled forms of Thele- 
phora caryophyllea but white; in Cuba 27. S. anastomosans 

12. Fructifications cespitose, somewhat creeping by tips of branches 

becoming attached to the matrix by disks; pileate branches 1-1 \ 
cm. long, 1-2 mm. broad; in Brazil 28. S. proliferum 

§IV. EFFUSO-REFLEXED SPECIES 

13. Hyaline, flexuous gloeocystidia conspicuous in the subhymenium and 

hymenium m 15 

13. Pyriform, vesicular organs present in trama, subhymenium, or hymenium . . . 16 

13. Colored conducting organs in trama, subhymenium, or hymenium; cystidia 

absent; hymenium bleeds when wounded, if in vegetative condition. 
S. hirsutum and S. rameale sometimes have occasional colored conducting 
organs in the hymenium 17 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



85 



17. 



13. Not having gloeocystidia, vesicular organs, nor colored conducting organs . . 14 

14. Hymenium lacking cystidia and paraphyses of noteworthy form or 

color :•%••• 1^ 

14. Cystidia present, incrusted or without incrustation and hair-like, 

hyaline, or colored 22 

14. Paraphyses noteworthy by bottle-brush form, branching, or color. 

For species having cystidia in addition to noteworthy paraphyses, 

see 27 . . . .. 28 

15. Coriaceous, dense, tawny, zonate, not sulcate, thin, 5-10 mm. in diameter; 

in Jamaica 29. S. caespitosum 

15. Soft, spongy, snuff-brown to bister, concentrically sulcate, reflexed 1-4 cm.; 

of wide range 80. S. fuscum 

15. Coriaceous-fleshy, bursting out from the bark, wart-like, peltate, vinaceous- 

brown, 2-4 mm. in diameter; no cystidia; on poplar. . .... .31. S. rufum 

15. Coriaceous-cartilaginous, shield-shaped, wood-brown, 1-4 mm. in diameter; 

cystidia present; on pine 82. S. Pint 

16. Coriaceous-soft, tomentose, lacking cystidia .83. S. purpureum 

16. Coriaceous-soft, tomentose, often with hairs becoming agglutinate 

into a rugose surface; hair-like cystidia present. .81+. S. rugosiusculum 
16. Corky, usually resupinate, sometimes reflexed and with the upper side 

a horny crust; vesicular bodies very numerous 35. S. Murrayi 

16. Stony hard throughout, the cut surface with a horn-like luster, 1-5 

mm. thick; vesicular bodies few; in Mexico and Jamaica . .36. S. saxitas 
Exuding a yellow milk, conducting organs of pale color; narrowly reflexed, 
tomentose; on Liquidambar and Carpinus in North Carolina and 

Alabama 87. S. styracifluum 

17. Milk red, conducting organs dark, numerous; fructifications cespitose- 

imbricated, villose to hirsute, tobacco-colored; on oak, Canada to 

Alabama and westward 38. S. gausapatum 

17. Milk red, conducting organs few; fructifications tomentose, concentrically 

sulcate, not cespitose; Florida to Brazil .39. S. australe 

17. Milk red, conducting organs dark, numerous; fructifications narrowly 

reflexed; hymenium multizonate; on frondose species, Newfoundland to 

North Carolina Jfi. S. rugosum 

Milk red, conducting organs numerous; on pine, spruce, and hemlock, 
Canada to Pennsylvania and westward to the Pacific coast. 41 . S. sanguinolentum 
18. Fructifications sulphur-colored, fading to cartridge-buff; inter- 
mediate layer not bordered by a golden, denser zone; Georgia to 

Brazil, and in Germany 42. S. sulphuratum 

18. Fructifications at first some shade of buff by reason of the hairy 

covering, becoming grayish with age, and at length often zonate and 
shining where disappearance of the hairy covering reveals the 

hardened, colored surface of the intermediate layer 19 

18. Fructifications white or whitish to cartridge-buff 20 

18. Fructifications snuff-brown or black above 21 

19. Effuso-reflexed, cream-buff at first, strigose-hirsute ; hymenium warm buff, 

sometimes pale smoke-gray; intermediate layer bordered by a narrow 
golden zone; colored conducting organs rarely present in the hymenium; 
Newfoundland to South Carolina and westward to the Pacific coast. 
43. S. hirsutum 

19. Effuso-reflexed at first, becoming umbonate-sessile, tomentose, sometimes 

with the tomentum becoming torn into narrow concentric bands rmd 
showing the bared surface chestnut in the furrows; margin not normally 
lobate; fructifications 2-7 cm. in diameter; common throughout North 
America # . . -44- S. fasciatum 

19. Wedge-shaped to umbonate-sessile, with a thinner covering of tomentum 

than S./asdaturrij becoming more bared and zonate than the latter, thinner 
and flexible, and with the margin normally cut into 2 or 3 large lobes; 
New York and Wisconsin southward to Brazil 45. 5. lobatum 

19. Covering of silky, villous fascicles arranged radially, becoming glabrous, 

shining, and radially ridged, not lobed nor folded together laterally, nor 
crisped; Florida to Dutch Guiana 46. S. versicolor 

19. Pilei 2-10 mm. long and broad, crowded together and folded or crisped, 

strigose-hairy towards the base; marginal portion shining and zoned, 



17. 



[Vol. 7 
86 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



cinnamon-buff to hazel; colored conducting organs occasionally present 

.••*. .••••. 47. S. rameale 

20. Fructifications 1-1 \ cm. in diameter, plane, thin, papery, silvery to 

pale gray and with a silky luster; common on Carpinus, Canada, 

eastern United States to Mexico 48. S. sericeum 

20. Fructifications 3-10 mm. in diameter, pubescent, white. 49. S. pubescens 
20. S. ochroleucum, an imperfectly known species of Europe, formerly 

reported in America, belongs here. For description of authentic 
specimen, see "species imperfectly known." 
20. Fructifications 2-4 mm. in diameter, conical, attached by the vertex 

and pendant, villose; Cuba 50. S. conicum 

21. Tobacco-colored, velvety-hirsute, becoming glabrous towards the margin 

and exposing the blackish, horny crust of the intermediate layer; hyme- 
nium pruinose; spores 4-5X2H*m; West Indies 51. S. vibrans 

21. Villose, blackening; intermediate layer not bordered by a crust; spores 

9X4/x; Mexico 52. S. crassum 

21. Velutinous and black above; coloring matter of intermediate layer dis- 
solved by KHO solution; hymenium ferruginous, radiately ridged; on 

coniferous wood, in northern United States 53. S. radiatum 

22. Cystidia hyaline, non-incrusted, hair-like 23 

22. Cystidia dark, or becoming dark, cylindric, obtuse, distinguishable 

from colored conducting organs by more or less granule-incrusta- 
tion; on conifers only 24 

22. Cystidia rough-walled or incrusted, somewhat colored either wholly 

or under the incrustation, pointed, not resembling conducting 
organs 25 

22. Cystidia incrusted, not at all colored except in S. cineraxcens at 

times; paraphyses not noteworthy 26 

22. Cystidia incrusted, not colored; paraphyses noteworthy by color or 

form 27 

23. Cinnamon to bone-brown, hoary; hair-like cystidia very few; spores 

(M0X3-4/*; Washington to New Mexico 64. S. patelliforme 

23. White, villose-tomentose; hymenium bright yellow; hair-like cystidia 

obtuse, 20-25 X4-6/z, numerous 55. S. ochraceo-flavum 

24. Coriaceous-spongy, dry, usually resupinate; hymenium pruinose, mul- 

tizonate; cystidia colored, cylindric, 90-150X6-8/*; spores 9-13 X 

4-5m ; northern United States 56. S. abietinum 

24. Very thick, felty, concentrically sulcate, drying with odor of anise; 

cystidia and basidiospores like those of S. abietinum; colored 
imbedded spores often present; Rocky Mountain states 

57. S. rugisporum 

24. Narrowly reflexed, tomentose, Prout's brown; hymenium umber; 

cystidia and spores as in S. abietinum; Vermont and New York. 

• • 58. S. ambiguum 

25. Vinaceous-lilac when young, becoming snuff-brown; cystidia colored, even, 

rough-walled or incrusted, 1 00-200 X 6-1 M ; from North Carolina and 

Ohio southward 59 . £. umbrinum 

25. Coriaceous-papery, thin, pliant, tomentose, concentrically sulcate, snuff- 
brown; hymenium velvety, snuff-brown, not multizonate; cystidia col- 
ored under the incrustation, conical, 30-75 Xl2-25/x; Florida to Brazil. 

• • 60. S. papyrinum 

25. With aspect of S. papyrinum but thinner; cystidia 45-60 X5-12m; hymen- 

ial layer 200/x thick; Jamaica 61. S. Earlei 

25. Coriaceous, tomentose, concentrically sulcate, hair-brown; cystidia slightly 

colored, roughened above, 50-120X4^,* on conifers, northern United 

States and Canada, and in Rocky Mountains 62. S. Chailletii 

25. Corky, usually resupinate, 1100/x thick; hymenium drab, 600m thick, con- 
taining innumerable, colored, rough cystidia 20-25 X5-7 M ; West Indies. 

••••■•■.••••♦••. 63. S. ferreum 

2b. btngose-hairy, concentrically sulcate, buff, weathering gray; hymen- 
ium pinkish buff to drab, bristling with cystidia 100-150 XI 2-20/u, 
sometimes brownish at the base; spores 10-12 X6/x. .64. S. cinerascens 
26. Coriaceous-gelatinous, small, whitish; cystidia 45-90 X12-15m; 

spores 15-20X12-14^; Jamaica 65. S. magnisporum 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 87 



26. Spongy-soft, tomentose; cystidia 36-60X9-12^; spores 5-9X3-4m; 

New York to Mexico 66. S. spumeum 

26. Bursting out from the bark, forming a gray hymenium and becoming 

reflexed, 1-2^ mm. in diameter .67. S. erumpens 

26. Corky, rigid, concentrically sulcate, bister; hymenium ruddy, be- 
coming zonate; cystidia 30-50X8-12/1/ spores 4-6X3-5m/ on 
hemlock and other conifers, Canada to Texas and westward to 
the Pacific coast 68. S. sulcatum 

26. Tobacco-colored and sulcate above, with a horn-like crust under the 

tomentum; hymenium whitish; cystidia 30-36 X7n; on oak, 

North Carolina and Ohio to Mexico 69. S. subpileatum 

27. With aspect of S. subpileatum as given above, but hymenium contains numer- 
ous and conspicuous bottle-brush paraphyses in addition to cystidia; 

Pennsylvania to Colombia 70. S. sepium 

27. With brown, velvety hymenium and a white margin; paraphyses filiform, 

colored; spores hyaline, 6-11X3-4^ 71. S. albobadium 

27. With aspect of dark specimens of S. albobadium, but with thicker, zonate 

hymenium and imbedded spores colored; Oregon to Mexico 

72. S. heterosporum 

27. Becoming narrowly reflexed, fuscous, 2-10 mm. in diameter; hymenium 

velvety, bister; paraphyses colored, with bushy-branched tips; Canada to 

Alabama and Arkansas 73. S. versiforme 

28. Snuff-brown and sulcate above, tomentose; hymenium pruinose, 

zoned, containing bottle-brush paraphyses; on oak, Florida and 
Venezuela 74* S. insigne 

28. Fuscous, sulcate, not tomentose but with upper surface a horn-like 

crust, 2-3 mm. thick; Mexico 75. S. durum 

28. Woody, resupinate, crowded as if confluent and then broken into 

frustules, 2-4 mm. in diameter, above black and crust-like; hyme- 
nium pinkish buff to whitish and pruinose ; on oak . . . .76. S. frustulosum 

28. Usually resupinate, coriaceous-soft; hymenium light vinaceous-purple 

when young, becoming avellaneous, containing filiform paraphyses 
with short lateral prongs; aspect of Corticzum evolvens; Canada to 
North Carolina 77. S. roseo-carneum 






i. Stereum caperatum (Berk. & Mont.) Massee, Linn. Soc. 
Bot. Jour. 27: 161. 1890; Lloyd, Myc. Writ. 4. Stip. Stere- 
ums, 17. textf. 531. 1913. Plate 2, fig. 1. 

Thelephora caperata Berkeley & Montagne, Ann. Sci. Nat. 
Bot. III. 11: 241. 1849; Montagne, Syll. Crypt. 175. 1856; 
Sacc. Syil. Fung. 6:523. 1888. 

Illustrations: Lloyd, loc. til.; Engl. & Prantl, Nat. Pflanzen- 



fam. (1: 1**): 124. f.H-J. 

Type: in Kew Herb. 

Pileus coracelous, infundibuliform, drying pinkish buff, the 
upper side with elevated radial ridges and usually densely tomen- 
tose with coarse fibers; in structure 600-700 n thick, composed of 
densely, longitudinally arranged, thick-walled, hyaline hyphae 
3 n in diameter; stem central or sometimes absent, with attach- 
ment by a tomentose disk; hymenium pale pinkish buff, some- 
what radially rugose, glabrous; hair-like cystidia not incrusted, 
3-4^ /* in diameter, flexuous, often constricted near the outer end, 



_ [Vol. 7 

88 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



protruding up to 12 n, are sometimes present; spores hyaline, 
even, 8-10 X 3-4 h M. 

Fructifications 2-10 cm. high, 2-15 cm. in diameter; stem, 
when present, 5 mm. - 2 cm. long, 2-5 mm. thick, often sessile. 

On decaying wood of frondose species. Florida, Louisiana, 
and West Indies to Bolivia. June to April, probably throughout 
the year. Common. 

S. caperatum is the largest infundibuliform Stereum of the 
Gulf states and the West Indies. Its large size, upper surface 
with elevated, radial ridges and usually heavy tomentum of 
coarse fibers, occurrence on wood to which it is attached by a 
villose or tomentose disk, constitute a group of characters by 
which the S. caperatum is readily recognized. Lloyd has pub- 
lished in his account of this species that it has true metuloids 
(incrusted cystidia) projecting 20-30 m, but I have found none 
whatever in either the type or in other collections referable to 
this species. 

Thelephora lamellataBerk. & Curtis, a species of Stereum related 
to S. caperatum and of rather similar aspect, occurring on islands 
of the Pacific, shows in the type specimen from Fiji Islands 
conical incrusted cystidia 6-12 n in diameter, protruding 12-25 m, 
and subglobose spores 3-3 \ X 3/». Since Lloyd cited S. caperatum 
as occurring in Samoa, the Philippines, and Australia, it is 
possible that his observations on incrusted cystidia of S. capera- 
tum were based on specimens from the Pacific region really 
referable to Stereum lamellatum rather than on the true S. 



from the Ame 



Hedwieia <a : 75 



1913, Bresadola gives T. lamellata as a synonym of Cladoderris 
infundibuliformis (Kl.) Fries. I have seen no American speci- 
mens referable to S. lamellatum. 

Specimens examined: 
Florida: New Smyrna, A. S. Bertolet; Ocala, W. H. Long, 12373 

(in Mo. Bot. Gard. Herb., 55125). 
Louisiana: A. B. Langlois, comm. by C. G. Lloyd, 2740; St. 

Martinville, A. B. Langlois, 2896 and an unnumbered 

specimen, C. J. Humphrey, 2518 (in Mo. Bot. Gard. 

Herb., 5111). 

Cuba: C. Wright, 290, 509 (in Kew Herb.); Candelaria, Earle 

& Wilson, 201; Guantanamo (in Weir Herb., 10858); 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 89 



Havana Province, P. Wilson, 1172, comm. by F. S. Earle; 

Herradura, Earle & Murrill, 180, comm. by N. Y. Bot. 

Gard. Herb. 
Porto Rico: Manati, Johnston & Stevenson, 2006 (in Mo. Bot. 

Gard. Herb., 3396). 
San Domingo: 259 (in Kew Herb.). 
Jamaica: Cinchona, L. M. Underwood, 3172 (in N. Y. Bot. 

Gard. Herb, and in Mo. Bot. Gard. Herb., 56271); Cockpit 

Country, E. G. Britton & D. W. Marble, 388 (in N. Y. Bot. 

Gard. Herb.). 
St. Kitts: Lambert Estate, N. L. Britton & J. F. Cowell, 672 

(in N. Y. Bot. Gard. Herb.). 
Brazil: Bahia, Blanchet, 19 (in Kew Herb.). 
Bolivia: Yungas, A.Miguel Bang, 295 (in Mo. Bot. Gard. Herb.). 



2. S. hydrophorum Berkeley, Ann. & Mag. Nat. Hist. I. 14: 
327. pi. 9. f. 2. 1844; Hooker's Jour. Bot. 8: 273. pi. 6. 
1856; Sacc. Syll. Fung. 6: 555. 1888; Massee, Linn. Soc. Bot. 
Jour. 27: 159. 1890; Lloyd, Myc. Writ. 4. Stip. Stereums, 
29. text f. 547, 548. 1913. Plate 2, fig. 2. 

Hymenochaete crater if or mis P. Hennings, Hedwigia 43: 172. 
1904; Sacc. Syll. Fung. 17: 166. 1905. 

Illustrations: Ann. & Mag. Nat. Hist. I. 14: pi. 9. f. 2; 
Hooker's Jour. Bot. 8 : pi. 6; Lloyd, loc. cit. 

Type: in Kew Herb. 

Pileus stipitate, coriaceous, infundibuliform, drying Prout's 
brown, obscurely zonate, velvety, sometimes bearing large, 

branched hairs at the center and bottom of the cups, the margin 
entire; stem central, cylindric, solid, velvety, colored like the 
pileus, enlarged at the base and attached by disk; hymenium 
even, drying snuff-brown, not setulose; in structure 600 n thick, 
composed of intermixed and interwoven hyaline and slightly 
colored hyphae, the latter of which give their color to the 
pileus and hymenium and curve into the hymenium as cylindric, 
obtuse, slightly colored paraphyses 3 n in diameter, not emergent 
above its surface; no cystidia, gloeocystidia, nor setae; spores 
hyaline, globose, even, 3 n in diameter. 

Pileus 4-10 cm. in diameter, 3-6 cm. deep; stem 3-5 cm. long, 
4-5 mm. thick. 



90 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



On wood on the ground. Venezuela, British Guiana, and 
Brazil. November. 

This South American species ranges so far to the north that it 
may possibly occur also in the West Indies or Central America. 
The fructifications have dimensions and general aspect of those 
of S. caperatum but are distinguishable by darker color of pileus, 
stem and hymenium, by velvety covering of pileus and stem, 

and by absence of elevated longitudinal ridges on the surface of 
the pileus. 

Specimens examined: 

Exsiccati: Ule, Myc. Brasil., 40, type distribution of Hymeno- 

chaete crateriformis. 
Venezuela: Maripa, M. A. Carriker, comm. by W. G. Farlow, 

III; Rio Mato, M. A. Carriker, comm. by W. G. Farlow, 

IV. 

Brazil: Spruce (in Curtis Herb.); Amazonas, Marmellos, E. 

Ule, in Ule, Myc. Brasil., 40. 



3. S. Ravenelii Berk. & Curtis, Grevillea i: 162. 1873; 

Sacc. Syll. Fung. 6: 552. 1888; Massee, Linn. Soc. Bot. Jour. 

27: 164. pi. 7. /. 2. 1890; Lloyd, Myc. Writ. 4. Stip. Stere- 

ums, 25. textf. 548. 1913. Plate 2, fig. 3. 

Illustrations: Lloyd, lac. cit.; Massee, loc. cit. 

Type: type distribution in Ravenel, Fungi Car. 4: 13. 

Fructifications gregarious, coriaceous, thin, often growing 

from a common mycelium; pileus infundibuliform, sometimes 

split on one side, even, drying cinnamon-buff 
to bay, often shining and zonate; stem slender, 
equal, minutely tomentose, drying pale olive-buff 
to pinkish buff; hymenium even, glabrous, colored 
like the stem; pileus in section 300-500 ^ thick, 
composed of densely and longitudinally arranged 
hyaline hyphae 3 m in diameter; flexuous gloeo- 
cystidia 30-60X4^-7 /x curve into the hymenium 
but do not protrude above its surface; no cysti- 



Fig. 1 . 



S. Ravenelii. dia; spores hyaline, even, 3-4X2^-3 m. 



Gloeocystidia X 

665. 



Fructifications 1^-5 cm. high, 3 mm-3 cm. 



From authentic in diameter; stem 5-10 mm. long, 



1 

2 



1* 



mm. 



specimen. 



thick. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 91 



On the ground, rarely on wood humus. South Carolina to 
Mexico, West Indies, and Brazil. July to April. 

S. Ravenelii is near S. pergamenum in microscopic characters 
but is constantly infundibuliform, with slender, more conspic- 
uous stem, and occurs on the ground except very rarely, and 
is gregarious rather than cespitose. The range of S. Ravenelii 
southward to Brazil is so much greater than has been noted 
heretofore that it would be well to compare with it authentic 
specimens of some of the imperfectly described South American 
species of central-stemmed Stereums 

Specimens examined: 
Exsiccati: Ravenel, Fungi Car. 4: 13. 
South Carolina: H. W. Ravenel, in Ravenel, Fungi Car. 4: 13, 

type distribution. 
Alabama: Beaumont, 207 in part (the small specimens on right 

of the card in Curtis Herb., 4629); Montgomery, R. P. 
Burke, 26, 181 (in Mo. Bot. Gard. Herb., 10305, 57059). 
Louisiana: Baton Rouge, C. W. Edgerton, 1544, and C. J". 

Humphrey & C. W. Edgerton, comm. by C. J. Humphrey, 
2523, 2522 (in Mo. Bot. Gard. Herb., 42921 and 42939 
respectively); St. Martinville, A. B. Langlois, 1847. 
Mexico: San Luis Potosi, C. G. Pringle (in Farlow Herb.). 
Cuba: C. Wright, 255 (under the name Stereum elegans in Curtis 

Herb.) ; Candelaria, Earle & Wilson, 205, 207; Herradura, 
N. L. Britton, E. G. Britton, F. S. Earle & C. S. Gager, 6397 
(in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 

56303). 

Trinidad: Carrengo, Carriker (in Farlow Herb., 1). 
Brazil: Blumenau, A. M oiler, the Stereum elegans of Hedwigia 

35: 288. 1896, comm. by G. Bresadola. 



4. S. surinamense Leveill£, Ann. Sci. Nat. Bot. III. 2: 209. 
1844; Sacc. Syll. Fung. 6:556. 1888; Massee, Linn. Soc. Bot. 
Jour. 27: 161. 1890; Lloyd, Myc. Writ. 4. Stip. Stereums, 26. 
textf. 5U- 1913. Plate 2, fig. 4. 

Stereum fulvo-nitens Berkeley, Ann. & Mag. Nat. Hist. II. 
9: 198. 1852; Sacc. Syll. Fung. 6:556. 1888; Massee, Linn. 
Soc. Bot. Jour. 27 : 162. 1890. 

Illustrations: Lloyd, loc. cit. 

Type: in Museum of Paris Herb, presumably. 



[Vol. 7 
92 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Pileus coriaceous, infundibuliform, sometimes more elongated 
on one side, glabrous, shining, lineate or striate, drying tawny to 
hazel, faintly zonate with numerous very narrow zones; stem 
central or eccentric, cylindric, drying avellaneous to burnt 
umber, fibrillose to minutely tomentose, attached at the base 
by a mycelial pad; hymenium glabrous, even, avellaneous to 
cinnamon; pileus in section 400 m thick, composed of a broad 
layer of densely and longitudinally arranged, thick-walled, 
hyaline hyphae 3 m in diameter and of a hymenial layer 45-90 m 
thick, the subhymenial portion of which may become thicker 
than the palisade layer of basidia and gloeocystidia and appears 
granular and composed of very fine hyphae; gloeocystidia 15-30 
m long, with ventricose base 6-9 m in diameter, sometimes barely 
emergent above the basidia; spores hyaline, even, 3-4X2-3 m. 

Fructifications 1^-4 cm. high, 1-2^ cm. in diameter; stem 

3-7 mm. long, about 1| mm. in diameter. 

On dead wood. West Indies, Honduras, and Dutch Guiana. 
November. 

Lloyd's account and figures have made possible the reference 
to S. surinamense of the collections cited below, for the original 
description by L6veille is fragmentary and does not even note 
whether the specimens were growing on the ground or on wood. 
I have not seen the types of either S. surinamense or S. fulvo- 
nitens. The specimens cited below are characterized by the 
attachment to the wood by a conspicuous mycelial pad, by 
rich hazel and shining upper surface of the large, narrowly 
zonate pileus, by the gloeocystidia, and by the minutely granular 
subhymenial region in which the hyphae are much finer than in 
the main hyphal layer and run at right angles to the latter. 

Specimens examined: 
San Domingo: Consuelo, N. Taylor, 176 (in N. Y. Bot. Gard. 

Herb, and Mo. Bot. Gard. Herb., 56293). 
Trinidad: R. Thaxter, comm. by W. G. Farlow (in Mo. Bot. 

Gard. Herb., 44304). 
British Honduras: M. E. Peck (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56326). 






5. S. macrorrhizum (L6veill6) Lloyd, Myc. Writ. 4. Stip. 
Stereums, 28. 1913. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 93 



Thelephora macrorrhiza LeVeill6, Ann. Sci. Nat. Bot. III. 
5: 146. 1846; Sacc. Syll. Fung. 6: 524. 1888. 

Type: in Museum of Paris Herb., according to LeVeille* and 

Lloyd. 

Pileus infundibuliform, coriaceous, somewhat membrana- 
ceous, rufescent, striatulate, the margin erect, possibly laciniate; 
hymenium sulcate, rather pallid; stem rather long, radicated. 

On ground, French Guiana. Coll. Melinon. 

Pileus coriaceous, nearly membranaceous, infundibuliform, 
russet, with rugosity from base to margin, the latter thin, lacini- 
ate; hymenium glabrous, rugose like upper surface of pileus; 
stem 1-2 decimeters long, glabrous, continued by a long radi- 
cated portion which extends perpendicularly into the ground. 
This character and also the absence of hairy covering of the stem 
afford a great difference between this species and Stereum 

surinamense. 

The above is a translation of the original description. I have 
not seen authentic specimens; Lloyd notes, loc. cit., that they 
are, "Stereum elegans, of an unusually regular growth. Not so 
confluent as ordinary." 



6. S. Burtianum Peck, N. Y. State Mus. Bui. 75: 21. pi 0. 
f. 30-34. 1904; Sacc. Syll. Fung. 17: 163. 1905; Lloyd, Myc. 
Writ. 4. Stip. Stereums, 21. text f. 537. 1913. Plate 2, fig. 5. 

Illustrations: Peck, loc. cit.; Lloyd, loc. cit. 

Type: in N. Y. State Mus. Herb, and in Burt Herb. 



Fructifications 



thin, infundibuliform 



sometimes split to the stem on one side, sometimes dimidiate, the 
upper surface slightly uneven with radiating fibrils and fibrillose 
ridges, cartridge-buff when fresh, drying Sayal-brown to hazel, 
the margin lobed or incised; stem solid, minutely tomentose, 
Sayal-brown in the herbarium; hymenium even or radiately 
uneven, glabrous, yellow ocher to pinard-yellow when fresh, 
becoming pinkish buff to Sayal-brown in the herbarium ; pileus 
in section 600 m thick, composed of densely and longitudinally 
arranged hyphae 2 m in diameter; no cystidia nor gloeocystidia; 
spores hyaline, even, subglobose, 3-4 m in diamter, or 4 X 3 m. 

Fructifications usually 12-20 mm. high, 5-15 mm. in diameter; 
stem 3-8 mm. long, 1-1 h mm. thick. 



n a [VoL - 7 

94 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



On the ground in frondose woods. New Hampshire to North 
Carolina and Tennessee, and in Japan. July to October. 

Distinguishing characters of this species are the radially 
arranged fibrils and fibrillose ridges of the upper surface of the 
pileus, bright yellow hymenium of fresh specimens, small sub- 
globose spores, and absence of zonation, cystidia, and gloeo- 
cystidia. These characters separate the species from S. auran- 
tiacum and S. Ravenelii and from specimens of S. diaphanum 
which have become discolored in the herbarium. The sections 
crush out and tissues spread apart when slight pressure is applied 
to the cover glass — a character unusual in stipitate Stereums. 
The specimen from Tennessee consists of two dimidiate pilei 
2X2^ cm. At Amherst, Massachusetts, Professor Anderson 
saw perhaps a thousand fructifications of this growing in an 
area of a square rod; to him I am indebted for the color obser- 
vations on fresh specimens and for specimens in growing condi- 
tion showing the colors and also the fact that the consistency 
of the pileus is not fleshy enough for inclusion of this species 
in Craterellus. 

Specimens examined: 
New Hampshire: Chocorua, W. G. Farlow, three collections 

(two of which are in Mo. Bot. Gard. Herb., 55242 and 55571, 

and the third in Farlow Herb.). 
Vermont: Lake Dunmore, W. G. Farlow (in Farlow Herb.). 
Massachusetts: Amherst, P. J. Anderson (in Mo. Bot. Gard. 

Herb., 56364, 56365). 
New York: Shokan, Ulster Co., C. H. Peck, type. 
North Carolina: Asheville, H. C. Beardslee, 2. 
Tennessee: Elkmont, C. H. Kauffman, 80 (in Mo. Bot. Gard. 

Herb., 44994). 
Japan: Sendai, A. Yasuda, 21 (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56290). 



7. S. rivulorum Berk. & Curtis, Linn. Soc. Bot. Jour 

io: 330. 1868; Sacc. Syll. Fung. 6: 552. 1888; Massee 

Linn. Soc. Bot. Jour. 27 : 167. 1890; Lloyd, Myc. Writ. 4. Stip 
Stereums, 21. 1913. 

Type: in Kew Herb, and probably in Curtis Herb. 

I failed to take any notes of the type specimens of this species 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 95 



when there was opportunity and have seen no collections which 
seem referable here. The translation of the original description 
follows : 

Minute, straw-colored; pileus cyathiform, decurrent into a 
stem dilated above, the margin undulate; hymenium glabrous. 

On wet ground amongst moss. Cuba, C. Wright, 533. 

Pileus 1| mm. across; stem 4 mm. high, oblique but not 
really lateral. Habit of a small stipitate Peziza. Spores glo- 
bose, 2-2^ m according to Massee. 



8. S. quisquiliare (Berk. & Curtis) Lloyd, Myc. Writ. 4. 
Stip. Stereums, 36. text f. 567. 1913. Plate 2, fig. 6. 

Thelephora quisquiliaris Berk. & Curtis, Linn. Soc. Bot. Jour. 
10: 329. 1868; Sacc. Syll. Fung. 6: 524. 1888. 

Illustrations: Lloyd, loc. cit. 

Type: in Kew Herb, and Curtis Herb. 

Pileus very small, flabellate or rarely cyathiform, tomentose, 
shining white; stem lateral, short, thickened above; pileus in 
section composed of loosely arranged hyphae 3-4 n in diameter; 
cystidia hair-like, not incrusted, 6 n in diameter, protruding up 
to 40 n beyond the basidia; spores hyaline, even, 4X3-4 n. 

Pileus 3-5 mm. broad, and 5-7 mm. long including the stem- 
like base. 

On particles of bark among moss and on mosses. Cuba. 

The fructifications of S. quisquiliare are small and of soft 
bibulous texture and resemble in aspect those of S. cyphelloides 
and Cyphella muscigena, but are distinguished from both these 
species by the hair-like cystidia, of which I noted the presence 



examination of the tvoe but which 



the permanent m 



I had hoped that 



would confirm the note as to hair-like cystidia 



and enable me to be more confident that Theleph 
should not be transferred to Cyphella. 

Specimens examined: 
Cuba: C. Wright, 519, type (in Curtis Herb.). 



9. S. aurantiacum (Pers.) Lloyd, Myc. Writ. 4. Stip. Stere 



urns, 22. text /. 538. 1913 



fig. 7 



Thelephora aurantiaca Persoon in Gaudichaud, Voy. Urania 
Bot. 176. 1827; Fries, Epicr. 536. 1838; R. Soc. Sci. Up- 



96 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



sal. Actis III. i: 108. 1851; Montagne in d'Orbigny, Voy. 
Am. Merid. Bot. 2: 48. 1839; in Ramon de la Sagra, Fl. 
Cub. 4: 228. pi 14. /• 1. 1853; Berkeley & Curtis, Linn. 
Soc. Bot. Jour. 10: 328. 1868; Sacc. Syll. Fung. 6: 526. 
1888. — T. sericella Berkeley & Curtis, Linn. Soc. Bot. Jour. 

— T. affinis 



1888. 



10: 328. 1868; Sacc. Syll. Fung. 6: 522. 
Berkeley & Curtis, Linn. Soc. Bot. Jour. 10: 329. 1868 (not 
T. affinis Pers.); Sacc. Syll. Fung. 6: 530. 1888. — Podoscypha 
aurantiaca (Pers.) Patouillard in Duss, Fl. Crypt. Antilles 
Fr. 230. 1904.— An T. spectabilis Leveille\ Ann. Sci. Nat. 
Bot. III. 2: 206. 1844? — An Stereum xanthellum Cooke, Gre- 
villeao: 12. 1880? 

Illustrations: Lloyd, loc. cit.; Montagne, loc. cit. 
Fructifications coriaceous, soft, everywhere drying Naples- 
yellow, losing the bright color in the herbarium; upper surface 

sericeous, lineate-striate, the margin variable, 
often somewhat fimbriate; stem thin, with yel- 
lowish tomentum at the base and sometimes with 
tomentose mycelial strands; hymenium even, or 

aline hairs under a 




nearly so, setulose with hy 



cystidia hair-like, not incrusted, cylindi 



obtuse 



diameter, protrudi 



spores hyaline, even, 5-8X3-4 n. 



Fructifications 2-3 



cm 



1-2 cm. 



Fig. 2. 

S. aurantiacum . 

Cystidium, basi- 

dia, and spores, 

X 665. 



in diameter when infundibuliform and 5 mm.-4 
cm. when flabelliform; stem 1 cm. long, about 
1 mm. thick. 

On ground and dead wood. West Indies to Par- 
aguay. June to February. Apparently frequent. 

S. aurantiacum is unique among the stipitate 
Stereums by its bright yellow color. Lloyd states 
that old specimens may lose their bright yellow 
color and become brown, and the figures by Mon- 



indicate this 



I have seen only one 



gathering in which some of the specimens have discolored brown 



ish; 



from Porto Rico, by Prof. Stevenson, bears 
rly pure white when collected: became vellow 



the field note : "nes 
in drying; no yellow showed until partly dried." The extensive 
synonymy of this species is due to its occurrence sometimes on 
the ground, sometimes on wood, sometimes being wholly infundi- 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 97 



buliform and sometimes wholly flabelliform, but occasionally 
a gathering shows both infundibuliform and flabelliform speci- 
mens. The soft texture of the pilei — like filter-paper or like 
wash leather — the large, cylindric, non-incrusted cystidia, and 
large elongated spores are a good combination of characters for 
the recognition of S. aurantiacum independently of the yellow 
color. Lloyd gives Thelephora spectabilis and Stereum xanthel- 
lum as synonyms of S. aurantiacum, and this seems quite prob- 
able according to the original descriptions of these species, but 
he does not state that he has studied the authentic specimens; 
I have not been able to examine them. 

Unless there is more than one edition of Gaudichaud's ' Voy. 
Urania Bot./ there is an error, as noted by Lloyd, in the citation 
by Fries in 'Epicrisis,' followed by later authors, of a figure of 
T. aurantiaca by Persoon. Dr. Farlow kindly searched for me 
for such a figure in his copy but without success. 

Specimens examined : 
Jamaica: Port Antonio, F. S. Earle, 600, comm. by N. Y. Bot. 

Gard. Herb.; A. E. Wight, comm. by W. G. Farlow; Troy 
and Tyre, W. A. Murrill & W. Harris, 1112, comm. by 
N. Y. Bot. Gard. Herb. 
Cuba: C. Wright, 237, type of Thelephora sericella (in Curtis 

Herb.); C. Wright, 198, 263, type of Thelephora affinis B. & 
C. (in Curtis Herb.); Banao Mts., Leon & Clement, 5570 
(in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 
56262) ; Ceballos, C. J. Humphrey, 2683 (in Mo. Bot. Gard. 
Herb., 8267); Guantanamo, Hioram (in J. R. Weir Herb., 
10583, and Mo. Bot. Gard. Herb., 56217); Omaja, C. J. 
Humphrey, 8025 (in Mo. Bot. Gard. Herb., 8632); Nipe 
Bay, F. S. Earle, No. A. 
Porto Rico: Rio Piedras, /. R. Johnston, comm. by J. A. Steven- 
son, 1987 (in Mo. Bot. Gard. Herb., 10660) ; /. A. Stevenson, 
3354, 5585 (in Mo. Bot. Gard. Herb., 17720 and 6908). 
San Domingo: Consuelo, N. Taylor, 178 (in N. Y. Bot. Gard. 

Herb, and Mo. Bot. Gard. Herb., 56304). 



io. S. diaphanum (Schw.) Cooke in Sacc. Syll. Fung. 6: 558. 
1888; Massee, Linn. Soc. Bot. Jour. 27: 162. 1890; Lloyd, 
Myc. Writ. 4. Stip. Stereums, 19. text f. 534. 1913. 

Plate 2, figs. 8 and 9. 
2 



[Vol. 7 



98 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Thelephora diaphana Schweinitz in Berk. & Curtis, Acad. Nat 

— T. Willeyi Clinton in Peck 



Sci. Phila. Jour 



278 



1853 



N. Y. State Mus. Rept. 26: 71. 1874; Sacc. Syll. Fung. 6:524 
1888.— An T. Sullivantii Montagne, Syll. Crypt. 176. 1856? 
Type : in Herb. Schweinitz, in Curtis Herb., and in Kew Herb 
Fructifications coriaceous, thin, deeply infundibuliform, some 
times deeply split, white, drying diaphanous, sericeous, fibril- 
lose, striate, sometimes with slij 




ridges, sometimes obscurely 



slightly elevated 
med, the margin 
thin, entire or laciniately toothed; stem slender, 
cylindric, more or less clothed with white matted 
down which is usually present at the base and 
binds the earth together in a ball ; pileus of type 



composed of longitud 



d 



walled hyaline hypl 



3 /i in 



diameter, densely 



ded together; hymenium 



Fig. 3. 

S. diaphanum. 

Cystidium, 

basidia, and 

spores, X 665. 



white, setulose with hyaline hairs under a lens; 
cystidia hair-like, not incrusted, cylindric, obtuse, 
6-9 m in diameter, protn 



dine 20-60 



n\ spores 



hyaline 



■5X21-3/* 



Fructifications 2-4 cm. high 



mm.-2 cm 



diamet 



mm. in diameter 



On the ground in moist 



ds of frond 



species 



New 



York to Missouri, and in Alabama, Washington, and California. 

S. diaphanum, as collected by Schweinitz and shown in pi. 

2, fig. 8, differs from S. aurantiacum in absence of bright yellow 

color, in shorter snores, and in stem and ground at base of stem 



being merely white-do 



New York 



attains a more luxuriant growth than the small specimens col- 
lected by Schweinitz, has a larger and rather thicker pileus and 



thicker stem as shown 
were published as The 



fig 



specimens 



Willeyi, but the intergradat 



with S. diaphanum are so numerous and close that it should be 
kept with the latter in my opinion. 

Specimens examined: 
New York: Buffalo, Clinton, type of Thelephora Willeyi (in N. 

Mus. Herb.); Chappaqua, Mrs. C. E. Ryder & 



Y. State Mus. 

Mrs, W. A. Murrill (in N. Y 



Bot. Gard. Herb, and Mo. 
Bot. Gard. Herb., 56289); Freeville, V. B. Walker, 15 






1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 99 



(in Mo. Bot. Gard. Herb., 8407) ; Geddes, G. E. Morris, G; 

Ithaca, C. Thorn (in Cornell Univ. Herb., 9992); Jamesville, 

H. D. House (in N. Y. State Mus. Herb, and in Mo. Bot. 

Gard. Herb., 55498), and L. M. Underwood; Lowville, 

C. H. Peck (in N. Y. State Mus. Herb.); Orville, G. E. 

Morris, G. 
Ohio: Gnaddenhutte, Schweinitz, type (in Herb. Schweinitz 

and in Curtis Herb.). 
Missouri: Valley Park, E. A. Burt & L. 0. Overholts (in Mo. 

Bot. Gard. Herb., 44059). 
Alabama: Montgomery, R. P. Burke, 25 (in Mo. Bot. Gard. 

Herb., 13146.). 
Washington: Seattle, W. A. Murrill, 128, 143, 1U (in N. Y. 

Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 55745, 55729, 

5572G). 

California: Tamalpais, H. W. Harkness (under the herbarium 

name Thelephora Harknessii Peck in N. Y. State Mus. 
Herb, and Mo. Bot. Gard. Herb., 55925). 



ii. S. exiguum (Peck) Burt, n. comb. Plate 2, fig. 10. 

Thelephora exigua Peck, N. Y. State Mus. Bui. 54: 953. 1902; 
Sacc. Syll. Fung. 17: 161. 1905. 

Type: in N. Y. State Mus. Herb, and in Burt Herb. 

Pileus coriaceous-membranaceous, very thin, diaphanous, 
infundibuliform, radiately fibrous-striate, becoming bister in the 
herbarium, originally "pale alutaceous" according to Peck, the 
margin lacerate; stem slender, solid, pruinose, and bearing a 
few whitish hairs which are present also on the ground about the 
base; pileus in section 100 /x thick, composed of longitudinally 
arranged, hyaline hyphae 2|-3 n in diameter, closely crowded 
together; cystidia hair-like, not incrusted, cylindric, obtuse, 
7 m in diameter, protruding 25 y. beyond the basidia; spores 
hyaline, even, 4|X2 n, borne 4 to a basidium. 

Fructifications 1-3 mm. in diameter, 3-5 mm. high; stem 2 
mm. long, \-\ mm. in diameter; pileus T V mm. thick. 

On the ground, Westport, New York. October. 

S. exiguum is miniature S. diaphanum of slightly darker 
color. It is known from the original collection only. The 
smallest specimens of S. diaphanum are many times larger than 



[Vol. 7 
100 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



the largest specimen of S. exiguum. While differences in size 

are not generally a good criterion for specific distinction, I am 
inclined to think that they will prove so in this instance. 
Specimens examined : 

New York: Westport, C. H. Peck, type (in N. Y. State Mus. 

Herb, and in Burt Herb.). 



12. S. tenerrimum Berk. & Ravenel, Grevillea i : 162. 1873; 
Sacc. Syll. Fung. 6: 551. 1888; Massee, Linn. Soc. Bot. Jour. 
27: 165. 1890. Plate 2, fig. 11. 

Type: in Kew Herb, and Curtis Herb. 

Pileus coriaceous, thin, infundibuliform or flabelliform, soon 
lobed and split, upper surface slightly rough, fibrillose-striate, 
not zonate or only very indistinctly "pale tan" when collected, 
becoming tawny olive to Saccardo's umber in the herbarium; 
stem filiform, whitish, bearing some fibrils towards the base; 
hymenium even, concolorous, setulose with hyaline hairs 
under a lens; pileus in section 300 n thick, composed of longi- 
tudinally and densely arranged hyaline hyphae 3 n in diameter ; 
cystidia hair-like, not incrusted, 4-8 m in diameter, protruding 
30-50 /x; spores hyaline, even, subglobose, 4-5 X 3-4 p. 

Fructifications 2-10 mm. broad, 5mm.-2| cm. high; stem 
3-7 mm. long, \-\ mm. thick. 

On ground among mosses. New York, Wisconsin, South 
Carolina, and Cuba. July to November. Rare. 

The collections which I have referred to S. tenerrimum are 
from the widely separated localities stated above and only a 
single gathering of several fructifications at each locality. There 
are slight differences between the specimens of the several 
gatherings, but not great enough to preclude their reference 
to a single species, although doing so has required some general- 
ization from the original description. 

S. tenerrimum is related to S. undulatum of northern Europe 
as known to me by the specimens distributed in Karsten, Fungi 
Fennicae, 912, and by the extended account by Maire, Ann. 
Myc. 7: 426-431, text /. 1, 2. 1909, but the latter species 
attains much larger size, has a coarser stem, and is infundibuli- 
form with central stem. None of the collections of S. tenerri- 
mum are composed wholly of specimens with infundibuliform 



1920] 



BURT — THELEPHORACEAB OF NORTH AMERICA. XII 



101 



pilei and the stem central; the original collections have some 
specimens with pileus longer on one side than the other and 
stem eccentric; in more recent gatherings some specimens are 
even flabelliform. S. tenerrimum appears to be a distinct 
species. 

Specimens examined: 
New York: Croghan, C. H. Peck (in N. Y. State Mus. Herb.). 
South Carolina: Society Hill, H. W. Ravenel, type (in Curtis 

Herb., 5029, and in Kew Herb.). 
Wisconsin: Afton, R. A. Harper. 
Cuba: Havana Province, Huo Leon, 1456 (in N. Y. Bot. Gard. 

Herb, and in Mo. Bot, Gard. Herb., 56307). 



13. S. pergamenum Berk. & Curtis, Gre\> 



161. 1873 



Sacc. Syll. Fung. 6 



1888; Massee, Linn. Soc. Bot. Jour 



27:161 



Lloyd, Myc. Writ. 4. Stip. Stereums 



545 



1913. 



/• 



Plate 2. fi 



An Stereum nitidulum Berkeley, Hooker's London Jour 



2: 638. 1843? 



distribution in Ravenel, Fund Car 



3: 25. 



Fructifications somewhat cespitose and grown together, stip- 
itate; pileus coriaceous, infundibuliform, sometimes split and 
petaloid, minutely lineate, drying hazel, obscurely 
zoned, the margin thin, often toothed or laciniate; 
stem cylindric, drying pinkish buff, very minutely 
tomentose; hymenium drying pinkish buff, gla- 
brous; pileus in section 500 n thick, composed 



of densely and longitudinally a 
hyphae 3 m in diameter; flexuous 



nged hv 



dia, 50 X 6 n, extend into the hymenium 



but do not rise to its surface; 



dia 



spores hyal 



even, slightly flattened on one 



side 



X3-3£ M 



Fructifications 1^-4 cm. hierh 



2 



mm.-3 cm 



diameter: stem 



mm 



diamet 



long, 1-3 mm. in 



Fig. 4. 

S. pergamenum. 
Gloeocystidia 



On stumps or buried wood, perhaps rarely 



X 665. 



on the ground. Ohio and North Carolina to Mexic 



d 



West 



September to January 



[Vol. 7 



102 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



S. pergamenum may be recognized by its occurrence in small 
clusters on wood at or near the surface of the ground, by small 
and nearly globose spores, and by the presence of gloeocystidia. 
It is probably more frequent in the West Indies than in the 
United States. When studying the specimens of this species 
in Kew Herbarium I compared with them the type of Stereum 
nitidulum Berk., collected by Gardner in Goyaz, Brazil, and 
concluded that it is probably the same species as S. pergamenum. 
In that early stage of my work I did not record the presence of 
gloeocystidia in the types of either S. pergamenum or S. nitidu- 
lum, and since I have no permanent preparation from the type 
of the latter, further, more critical study may show that it is a 
distinct species. The collection from Cuba, referred by Ber- 
keley to S. nitidulum, has gloeocystidia and is referable to S. 

pergamenum. 

Specimens examined: 
Exsiccati : Ravenel, Fungi Car. 3 : 25. 
Ohio: Preston, T. G. Gentry (in N. Y. Bot. Gard. Herb, and Mo. 

Bot. Gard. Herb., 56301). 
North Carolina: Blowing Rock, G. F. Atkinson, from Bot. Dept. 

of Cornell Univ., 4182. 

Alabama: J. M. Peters, in Ravenel, Fungi Car. 3: 25, type 

distribution; J. M. Peters, 601 and another specimen (in 
Curtis Herb., the latter, Curtis Herb., 3814); Beaumont, 
207 in part, the large zonate specimen mounted on left side 
of card with specimens of S. Ravenelii (in Curtis Herb., 
4629 in part); Auburn, F. S. Earle (in N. Y. Bot. Gard. 
Herb, and Mo. Bot. Gard. Herb., 56306). 

Louisiana: Ville Platte, A. B. Langlois, 2897. 

Mexico: Motzorongo, near Cordoba, W. A. & Edna L. Murrill, 

994 (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 

54596). 
Cuba: C. Wright, 836 (in Curtis Herb., under the name S. 

nitidulum Berk.); Herradura, F. S. Earle, 5 45, and N. L. 
Britton, E. G. Britton, F. S. Earle & C. S. Gager, 6326 (in 
N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 56305 
and 56263 respectively); Sumidero, J. A. Shafer, 13905 
(in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 
56264). 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



103 



San Domingo: Bonao, J. A. Stevenson, 7010 (in Mo. Bot. Gard. 

Herb., 55656). 






14. S. cristatum Berk & Curtis, Grevillea 1: 163. 1873; 
Sacc. Syll. Fung. 6:556. 1888; Massee, Linn. Soc. Bot. Jour. 
27: 167. 1890; Lloyd, Myc. Writ. 4. Stip. Stereums, 38. 
1913. 

Type: in Kew Herb., not found by me in Curtis Herb, al- 
though sought for. 

Pileus coriaceous, flabelliform or obliquely cyathiform, pallid 
to light bay-brown, somewhat zoned, glabrous and shining 
towards the margin, bearing a cluster of coarse 
hairs towards the base; stem, when present, 
cylindric, scarcely 2 mm. long; hymenium even, 
paler than the upper surface ; in structure 200-250 
/x thick, composed of longitudinally arranged and 
somewhat interwoven hyaline hyphae 3 /* in 
diameter; no cystidia; gloeocystidia pyriform, 
9-12X7^ n) spores, as found in a crushed prepara- 
tion, hyaline, even, 4X2^ ^ few found — noted by 
Massee as subglobose, 5-6 /x in diameter. 

Pileus 6-10 mm. across. 

On dead Vitis in swamps. South Carolina. 

Reexamination of my preparation of the type 




Fig. 5. 

S. cristatum. 
Gloeocystidia X 



of S. cristatum fails to demonstrate that the 665. From type. 
pyriform organs in its hymenium are longitu- 
dinally septate; furthermore some of these organs are more 
elongated than stated above and irregular in form. For 
these reasons I regard the bodies as pyriform gloeocystidia 
rather than possibly miniature basidia of the longitudinally 
septate type, the demonstrated presence of which would require 
transfer of this species to Eichleriella. The occurrence of S. 
cristatum on dead grape vines, the crest of coarse hairs towards 
the base of the pileus, the small size of the latter, and the pyri- 
form organs in the hymenium are a good group of characters 
for identification of this species, although known so far only 
from the original collections. 

Specimens examined: 
South Carolina: Santee Swamp, H. W. Ravenel, Curtis Herb. 






104 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



No. 2038, type and an unnumbered specimen (both in Kew 
Herb.). 



15. S . pallidum ( Pers . ) Lloyd, Myc. Writ. 4. Stip. Stereums, 



31. textf. 536, 550. 



1913. 



Plate 3, fig. 13, 14. 



Craterella pallida Persoon, Ic. et Descr. Fung, i: 3. pi. 1. 




3. 



1798. 



Thelephora pallida Persoon, Syn. Fung. 565. 



1801; Myc. Eur. i: 111. 1822; Fries, Hym. Eur. 633. 



1874; Sacc. Syll. Fung. 6: 527. 



1888. 



Helvella 



pannosa 



Sowerby, Col. Figs. Eng. Fungi, pi 155. 1788, in part. — Thele- 
phora pannosa Sowerby ex Fries, in part, and T. pannosa var. 



pallida (Pers.) Fries, Syst. Myc. i : 430. 



1821. 



T. Sowerbeyi 



Berkeley, Outlines Brit, Fungi, 266. 1860; Ann. & Mag. Nat. 
Hist. III. 15:320. 1865; Fries, Hym. Eur. 633. 1874; Sacc. 
Syll. Fung. 6: 522. 1888. — Stereum Sowerbeyi (Berk.) Massee, 
Linn. Soc. Bot. Jour. 27: 164. 1890. — Bresadolina pallida 
(Pers.) Brinkmann, Ann. Myc. 7:289. 



1909. 

Illustrations: Persoon, Ic. et Descr. Fung. i:pl. 1. f. 3; 
Sowerby, Col. Figs. Eng. Fungi, pi. 155; Lloyd, Myc. Writ. 

4. Stip. Stereums, textf. 536, 550. 

Fructifications cespitose, hit orally confluent, 
infundibuliform, coriaceous-spongy, rather thick, 
becoming cartridge-buff to cream-color in the 
herbarium, the upper side strigose-squamose; 
stem short, villose at the base; hymenium with 
slight, very obtuse, radial folds, under a lens 
more or less setulose with hyaline hairs; cystidia 
hair-like, not incrusted, cylindric, 6-8 m in di- 
ameter, protruding 10-50 n beyond the basidia, 
usually very numerous but sometimes only few 
found; spores hyaline, even, flattened on one 
side, 6-8X4-5 /z. 

Fructifications 1-3 cm. in diameter, 2-3 cm. high. 

On the ground in woods. Vermont to North 
Carolina. July to November. Rare. 

American specimens of S. pallidum agree 




Fig. 6. 

S. pallidum. 



Cystidmm, basi- we jj w itli the European specimen received from 

x 665. From' Bresadola, and, like the latter, are paler than 
Bresadola. the otherwise excellent figures of Thelephora 



19201 in* 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 1U5 



pallida in Persoon's 'Icones et Descriptiones Fungorum' already- 
cited. Our specimens and that from Bresadola have the 
hymenium distinctly setulose with hair-like cystidia. Some of 
the specimens in Kew Herbarium under the name of Thelephora 
Sowerbeyi have hair-like cystidia, but these organs are few or 
absent in whole sections from other specimens. The original 
specimen of Helvella pannosa from Sowerby in Berkeley Her- 
barium at Kew has hair-like cystidia. I concluded that these 
cystidia are variable in abundance in English specimens and 
that Thelephora Sowerbeyi and Helvella pannosa as represented 
by the specimen from Sowerby should be kept with Thelephora 
pallida. Although the specific name pannosa of Sowerby was 
at first adopted by Fries, this was dropped later when Berkeley 
found this species, as understood by Sowerby, to be based upon 
a mixture of two species which were separated as Thelephora 
Sowerbeyi and T. multizonata ; T. pallida has priority over T. 

Sowerbeyi. 

S. pallidum may be distinguished from T. Willeyi forms of S. 
diaphanum by its occurrence in small concrescent clusters, by 
short villose or tomentose stem, and by thicker pileus with upper 
surface split radially into stiff straight fibrils. 

Specimens examined: 
Austria: G. Bresadola. 
England: from Sowerby, under the name Helvella pannosa (in 

Kew Herb.); Cornwall, C. Rea, 1 (in Mo. Bot. Gard. Herb., 

56241); Hereford, Mrs. Wynne (in Kew Herb., under the 

name Thelephora Sowerbeyi). 
Vermont: Brattleboro, C. C. Frost (in Univ. Vermont Herb.); 

Grand View Mountain, E. A. Burt. 
Connecticut: Waterbury, C. C. Hanmer, 1191. 
North Carolina: Blowing Rock, G. F. Atkinson, comm. by 

Cornell Univ. Herb., 4192. 

16. S. elegans (Meyer) Lloyd, Myc. Writ. 4. Stip. Stereums, 
24. textf. 589. 1913. (Not S. elegans of earlier authors.) 

Plate 3, fig. 15. 

Thelephora elegans Meyer, Fl. Essequeboensis, 305. 1818; 
Fries, Syst. Myc. 1 : 430. 1821; Epicr. 545. 1838. (But here 
abridged in an important respect so that following authors 
modified the description to apply to more common species). 






106 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 




An T. macrorrhiza LeVeille, Ann. Sci. Nat. Bot. III. 5: 146. 
1818? See Lloyd, loc. cit., p. 28. 

Illustrations: Lloyd, loc. cit. Not by the figures under this 
name in other works, as Engl. & Prantl, Nat. Pflanzenfam., for 
example. 

Fructifications cespitose, coriaceous, confluent, infundibuli- 
form and deeply split on one side, or little developed on one side 

and prolonged and petaloid on the other; upper 

surface of pilei glabrous, radially plicate, drying 

diamine-brown, the margin paler and more or less 
lobed; stems solid, buffy brown, short, tomen- 
tose, branched above ; hymenium radially plicate, 
nearly white, pruinose, often cracked; pileus in 
section 400 m thick, composed of densely and longi- 
tudinally arranged, hyaline hyphae 3 n in diam- 
eter; no cystidia; gloeocystidia 4| n in diameter, 
barely distinguishable from the basidia; spores 
hyaline, even, subglobose, 3f-4§ m in diameter. 

Fructifications 4-5 cm. high; pilei 1-2 cm. in 
diameter; stems about 1 cm. long, 1-2 mm. in 
diameter. 

In a dense cluster of about 16 fructifications 
springing from an area of 2 square centimeters 
on the ground. Porto Rico to British Guiana. Summer. 

I have not seen the type of Stereum elegans from Dutch Guiana 
nor reference to its existence; a collection from Porto Rico on 
which the preceding description is based has fructifications 
growing on the ground closely together and concrescent where 
in contact; the pilei are plicate on both surfaces and contrast 
so greatly in color that it seems as though fuscous in connection 
with the upper side and whitish flesh-color and pruinose for the 
under side might have been used for the color difference. The 
specimens of this collection are not zonate; infundibuliform 
without any qualification of this character does not seem accu- 
rate ; hence it may be that this Porto Rican collection is merely 
near, rather than the true, Stereum elegans. However, solitary 
fructifications growing on wood, as figured in Engl. & Prantl, 
Pflanzenfam., are certainly a very different species from S. 
elegans, the original description of which is as follows: 



Fig. 7. 

S. elegans. 

Gloeocystidia 

X 665. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 1U7 



<< 



1. Thelephora elegans. nob. 
"T. subcaespitosa infundibuliformis carnoso-coriacea plicata 
utrinque glabra, superne dilute fusco-fasciata, inferne albescenti- 

carnea pruinosa. 

"Ad terram argillosam. 

"Viget Junio. 

"Adumbr. Pulchra species. Gregarie crescens, subcarnosa, 
tenuis, glabra. Pileus substipitatus, 1-2 uncialis, infundibuli- 
formis, subcompressus, undulato-plicatus, margine irregulariter 
crenatus, interne rufescens, et fasciis dilute fuscis eleganter vari- 
egatus, nitens, externe albescenti-carneus, opacus, pruinosus 

Specimens examined: 
Porto Rico: Mayaguez, B. Lopez Santiago, 17 (in N. Y. Bot. 

Gard. Herb, and Mo. Bot. Gard. Herb., 56265). 



» 



7. S. decolorans (Berk. & Curtis) Lloyd, Myc. Writ. 4 






Stip. Stereums, 36. 1913. Plate 3, fig. 234. 

Thelephora decolorans Berkeley & Curtis, Linn. Soc. Bot. Jour. 

10: 328. 1868; Sacc. Syll. Fung. 6: 530. 1888.— Podoscypha 

decolorans (Berk. & Curtis) Patouillard in Duss, Fl. Crypt. 

Antilles Fr. 231. 1904. 

Type: in Kew Herb, and Curtis Herb. 

Fructifications coriaceous, gregarious or somewhat cespitose, 
stipitate; pileus split on one side quite, or nearly, to the stem, 
usually wedge-shaped to broadly flabelliform, sometimes radially 
lineate, drying cinnamon; stem cylindric, colored like the 
pileus, tomentose, attached by a mycelium common to several 
fructifications; hymenium colored like the pileus and stem, 
sometimes lineate ; pileus in section 200-400 /* thick, composed 
of densely and longitudinally arranged hyaline hyphae 3-3£ m 
in diameter; no cystidia; gloeocystidia flexuous, 45-90X3-6 m, 
between the basidia or curving into the hymenium; spores 
hyaline, even, subglobose, 4-4^X3-4 m. 

Fructifications 1-3 cm. long, 5-13 mm. broad; stem 2-10 



mm 



mm. thick 



On dead wood. Jamaica to Trinidad. May to January. 

S. decolorans is stated in the original description to have been 
white, drying ochraceous; I have seen only dried specimens 
which are pale cinnamon throughout. The occurrence of the 



1 C\Q . lVoL - 7 

1U8 ANNALS OF THE MISSOURI BOTANICAL GARDEN 

fan-shaped fructifications in clusters on dead wood, pale cinna- 
mon color when dry, presence of gloeocystidia, and small sub- 
globose spores constitute a group of characters by which dried 
specimens of S. decolorans may be distinguished from other 
species in our region. 

Specimens examined: 
Jamaica: W. A. Murrill, 1181 (in N. Y. Bot. Gard. Herb.). 
Cuba: C. Wright 234, %48, type (in Kew Herb, and Curtis 

Herb.); Santiago de las Vegas, Van Herman, comm. by 

F. S. Earle, 257. 

Trinidad: Carengo, M. A. Carriker, comm. by W. G. Farlow, 1. 

18. S. radicans (Berk.) Burt, n. comb. Plate 3, fig. 16. 

Thelephora radicans Berkeley, Hooker's London Jour. Bot. 
3: 190. 1844; Berkeley & Curtis, Linn. Soc. Bot. Jour. 10: 
329. 1868; Sacc. Syll. Fung. 6: 525. 1888.— Podoscypha radi- 
cans (Berk. & Curtis) Patouillard in Duss, Fl. Crypt. Antilles 
Fr. 230. 1904. 

Type: in Kew Herb, probably. 

"Plant 1£ inch high, f of an inch broad, spathulate or subin- 
fundibuliform, split on one side and slightly lobed, minutely 

lines, tawny, coriaceous. Stem 1 of an inch 



d 



high, 1± line thick, incrassated at the base, and sending off 
strong branched roots. Hymenium nearly even, fuliginous; 
spores apparently fuliginous." 

The above is the original description of the type specimens, 
collected in Surinam, Guiana, by Hostmann, 489. My knowl- 
edge of the species is based upon a later collection made in Cuba 
by C. Wright and determined by Berkeley. This specimen 
and the others cited below show well the longitudinal raised 
lines on the upper surface of the pileus, which is thicker than in 
related species, being 1-1 \ mm. thick, and the hymenium 100- 
200 n thick; some specimens have dried with the upper surface 
pinkish buff and others from wood-brown to Verona-brown: 
hymenium even, wood-brown to fuscous; stem 10-15 
3-4 mm. in diameter, sometimes radicated to reach buried wood; 
no cystidia nor gloeocystidia; spores hyaline, even, becoming 
minutely rough-walled and sometimes slightly angular, 6 X 5 p. 

Specimens examined: 
Cuba: C. Wright, 209, authentic (in Curtis Herb.). 



mm 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 109 



Jamaica: Castleton Gardens, W. A. & Edna L. Murrill, 66 > 

comm. by N. Y. Bot. Gard. Herb. 
Trinidad: R. Thaxter (in Farlow Herb.). 
Grenada: W. E. Broadway, September collection (in N. Y. Bot. 

Gard. Herb, and Mo. Bot. Gard. Herb., 5(3316) ; St. George's, 

W. E. Broadway (in N. Y. Bot. Gard. Herb, and Mo. Bot. 

Gard. Herb., 56317). 
British Honduras: M. E. Peck (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56321). 

19. S. pusiolum Berk. & Curtis, Linn. Soc. Bot. Jour. 10: 330. 
1868; Sacc. Syll. Fung. 6: 558. 1888; Massee, Linn. Soc. Bot. 
Jour. 27: 168. 1890; Lloyd, Myc. Writ. 4: Stip. Stereums, 39. 
1913. Plate 3, fig. 17. 



Type: in Kew Herb, and Curtis Herb. 

Fructifications gregarious, stipitate, coriaceous, curling in 
drying; pileus flabelliform or wedge-shaped, tapering to the 
stem, more or less split when large, minutely tomentose or 
hoary, white at first, drying smoke-gray, the margin thick and 
entire; stem short, solid, a little larger towards the base, colored 
like the pileus; hymenium even, mouse-gray, thick, contracting 
and sometimes cracking in drying; pileus in section 400-800 
n thick, composed of closely and longitudinally arranged hyaline 
hyphae 2^ in diameter; no cystidia, gloeocystidia, nor conduct- 
ing hyphae; spores hyaline, even, apiculate at base, 4-5| X 3-5 n. 

Fructifications 1-2 cm. high, 1-15 mm. broad; stem 5-8 mm. 
long, \-\\ mm. thick. 

On clay ground. West Indies. November to March. 

The white pileus, drying gray of nearly the shade of Polyporus 
adustus, minutely hairy, wedge-shaped, and without zonation, 
the much darker hymenium — dark as in P. adustus — the rather 
large spores, and the absence of gloeocystidia afford a group of 
characters highly distinctive for Stereum pusiolum, the descrip- 
tion of which I have changed materially from that published 
by the authors of the species. They disregarded Wright's note 
that the specimens were white and were collected on banks by 
roadside and published instead " rufobrunneum" and "on root- 
lets. " The recent collections, cited below, which I have com- 
pared with the type, show also that the dimensions of the fructi- 
fications are usually much larger than those of the type collection. 



[Vol. 7 
110 ANNALS OF THE MISSOURI BOTANICAL GARDEN 






Specimens examined: 
Cuba: C. Wright, 510, type (in Curtis Herb.); El Yunque, 

Baracoa, L. M. Underwood & F. S. Earle, 1087, 11 41, 

comm. by N. Y. Bot. Gard. Herb., 1141 (in Mo. Bot. Gard. 

Herb., 56588). 
Porto Rico: Rio Piedras, J. R. Johnston, 89 (in N. Y. Bot. 

Gard. Herb, and Mo. Bot. Gard. Herb., 56284). 

20. S. glabrescens Berk. & Curtis, Linn. Soc. Bot. Jour. 
io : 330. 1868; Sacc. Syll. Fung. 6: 558. 1888; Massee, Linn. 
Soc. Bot. Jour. 27: 169. 1890; Lloyd, Myc. Writ. 4. Stip. 
Stereums, 37. text f. 558. 1913. Plate 3, fig. 18. 

Illustrations: Lloyd, loc. cit. 

Type: in Kew Herb, and Curtis Herb. 

Fructifications scattered, sometimes two from a common 
mycelial pad, stipitate; pileus flabelliform, zonate, minutely 
velvety, sometimes nearly glabrous, drying Verona-brown to 
chestnut, the margin paler, tapering behind into a short stem; 
stem lateral, nearly equal, velvety; hymenium even, concave, 
drying pinkish buff; no cystidia nor gloeocystidia; spores 
hyaline, even, 4-5X3-4 n. 

Pileus 5-20 mm. long, 5-20 mm. broad; stem 2-10 mm. long, 
5-I3 mm. thick. 

On fallen twigs and mossy rotten wood. West Indies. May 
to September. 

S. glabrescens has small, rather scattered fructifications, with 
firm, coriaceous, minutely velvety pileus and stem, small sub- 
globose spores, and no cystidia, and it occurs on wood. Some 
collections are nearly glabrous. A mycelial pad is usually 
present at base of stem. 

Specimens examined: 
Cuba: C. Wright, 520, type (in Curtis Herb.); Pinar del Rio, 

J. A. Shafer, 13906 (in N. Y. Bot. Gard. Herb, and Mo. 
Bot. Gard. Herb., 56298). 
Porto Rico: Ponce, F. S. Earle, 163, comm. by N. Y. Bot. Gard. 

Herb. 
Jamaica: Hollymount, L. M. Underwood, 3427 (in N. Y. Bot. 

Gard. Herb, and Mo. Bot. Gard. Herb., 56299). 
Dominica: Landat, F. E. Lloyd, 380, comm. by N. Y. Bot. 

Gard. Herb. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 111 



21. S. flabellatum Patouillard, Soc. Myc. Fr. Bui. 16: 179. 
1900; Sacc. Syll. Fung. 16: 187. 1902; Lloyd, Myc. Writ. 
4. Stip. Stereums, 39. 1913. 

Podoscypha flabellata Patouillard in Duss, Fl. Crypt. Antilles 
Fr. 231. 1904. 

Pileus membranaceous, thin, expanded anteriorly, regularly 
attenuated posteriorly into a lateral stipe which is compressed ; 
margin papyraceous, deeply incised or lobed; dorsal surface 
marked by slight puberulence of projecting hairs or crests which 
are slightly diverging or fan-shaped, not zonate; hymenium 
inferior, glabrous, even; stem becoming pubescent, short, 
enlarged at the base into a disk for attachment. 

Fructification 4-6 cm. high; stem §— 1 cm. long, 1-2 mm. 
thick. Fructification erect, spathulate, often confluent by the 
margin with neighbors, whitish when living, livid and pellucid 
upon drying. 

On rotting wood on the ground. Guadaloupe. 

The above is a translation of Patouillard's description. Lloyd 
saw a specimen in the museum at Berlin and states that the 
dried specimens are dark reddish bay. 



22. S. fissum Berkeley, Hooker's Jour. Bot. 8: 273. 1856; 
Massee, Linn. Soc. Bot. Jour. 27: 169. 1890; Sacc. Syll. 
Fung. 11: 120. 1895; Lloyd, Myc. Writ. 4. Stip. Stereums, 
37. text f. 559. 1913. Plate 3, fig. 19. 

S. H uberianum P '. Hennings, Hedwigia 41: (15). 1902; 43: 
173. 1904. 

Illustrations: Lloyd, loc. cit. 

Type: in Kew Herb, and in Curtis Herb. 

Pilei gregarious, occurring singly, sessile or short-stipitate, 
coriaceous, flabelliform or wedge-shaped, often divided into 
wedge-shaped segments, glabrous, even, not shining nor zonate, 
white when fresh, now reddish brown in the herbarium, attached 
by a flat mycelial pad; hymenium even; in structure 300-400 
n thick, composed of densely and longitudinally arranged hyaline 
hyphae 3 n, or some 4 ju, in diameter; no cystidia nor gloeo- 
cystidia; the few detached spores found are hyaline, even, 6X4 ^. 

Pileus 8-15 mm. long, 3-15 mm. broad. 

On dead twigs, Brazil. 



(Vol. 7 
112 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



S. fissum may yet be found as far north as the West Indies and 
Central America. The species is noteworthy by its occurrence 
on dead twigs in scattered, solitary, azonate fructifications which 
are often deeply split into segments, and by absence of cystidia 
and gloeocystidia. 

Specimens examined: 
Exsiccati: Ule, Myc. Brasil., 42, under the name Stereum Huberi- 

anum. 
Brazil: Panure, Spruce, 27, type (in Curtis Herb.); Amazonas, 

Marmellos, and Jurnd, E. Ule, in Ule, Myc. Brasil., 42. 



23. S. cyphelloides Berk. & Curtis, Linn. Soc. Bot. Jour. 10: 
331. 1868; Sacc. Syll. Fung. 6: 558. 1888; Massee, Linn. 
Soc. Bot. Jour. 27: 172. 1890; Lloyd, Myc. Writ. 4. Stip. 
Stereums, 35. 1913. Plate 3, fig. 20. 

Type: in Kew Herb, and Curtis Herb. 

Pileus small, flabelliform or spatulate, drying pinkish buff, 

longitudinally fibrillose, bibulous, the margin entire, narrowed 
behind into a short stem-like base; in structure up to 600 /* 
thick, composed of thin-walled, hyaline hyphae 2^-3 n in diam- 
eter, interwoven in the subhymenium; hymenium even, dry- 
ing of same color as upper surface of pileus; no conducting 

organs, gloeocystidia, nor cystidia; spores hyaline, even, 4-5 X 
3-3 \ ix. 

Pileus 3-6 mm. wide, 5-7 mm. long. 

On a bank among moss. West Indies. February and March. 

S. cyphelloides differs from most Stereums in not having a hard 
compact structure, as in S. rameale, for example; it is of soft and 
bibulous texture but rather too thick for a Cyphella. The stem- 
like base is flattened in the same plane with the pileus and has the 
hymenium continued along its whole length, hence it is merely a 
narrowed portion of the pileus. 

- 

Specimens examined: 
Cuba: C. Wright, 511, type (in Curtis Herb.). 
Porto Rico: Monte Cerrote, near Adjuntas, N. L. Britton & 

Stewardson Brown, 5449 (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56261). 



24. S. Hartmanni (Mont.) Lloyd, Myc. Writ. 4. Stip. Stere- 
ums, 34. text f. 553. 1913. Plate 3, fig. 21. 



1920] 

BURT— THELEPHORACEAE OF NORTH AMERICA. XII 113 



gments 



Thelephora Hartmanni Montagne, Ann. Sci. Nat. Bot. II 
20: 366. 1843; Syll. Crypt. 176. 1856; Sacc. Syll. Fung. 6 
535. 1888.— T. dissecta Leveille, Ann. Sci. Nat. Bot. III. 5 
146. 1846; Sacc. Syll. Fung. 6: 531. 1888; Lloyd, loc. tit., 39 

Illustrations: Lloyd, loc. tit. 

Type: authentic specimen from Montagne in Kew Herb. 

Pilei solitary or cespitose, sessile or barely stipitate, coriaceous 
thin, white, wedge-shaped, deeply cleft into narrow si 
which are more or less pectinate along their margins and apex 
and have these teeth-like portions incurved; no cystidia; no 
gloeocystidia; spores hyaline, even, subglobose, 4-5X3^-4 y.. 

Pileus 7-50 mm. long, 5-40 mm. broad. 

On decaying wood and bark and dead herbaceous stems. 
Carolina to Bolivia. July to September in West Indies and 
February in Bolivia. 

The pilei of S. Hartmanni occur in small tufts of two or three 
in the specimens which have been seen; they are very dainty 
and unique by the narrow pectinate margins and tips which are 
more or less incurved; rarely these teeth occur on the lower 
surface of segments of the pileus in a manner suggestive of teeth 
of an Irpex but they are in most cases marginal. The maximum 
dimensions of the pileus are from the Porto Rican collection; 
the other specimens do not have pilei more than 2-3 cm. long. 
I have not seen the type of Thelephora dissecta Lev., which was 
collected in Guadeloupe; the description agrees so well with S. 
Hartmanni that I have followed Lloyd's conclusion that T. 
dissecta is a synonym of S. Hartmanni. 

Specimens examined: 
Carolina: Hartmann, authentic, from Montagne (in Kew Herb.). 
Porto Rico: Luquillo Mountain, P. Wilson, SIS (in N. Y. Bot. 

Gard. Herb, and Mo. Bot. Gard. Herb., 56302). 
St. Kitt's: N. L. Britton & J. F. Cowell, 706, comm. by N. Y. 

Bot. Gard. Herb. 

Bolivia: R. E. Fries, 272, comm. by L. Romell, 447 (in Mo. 

Bot. Gard. Herb.. 54780). 



25. S. craspedium (Fries) Burt, n. comb. Plate 3, fig. 22. 

Thelephora (Merisma) craspedia Fries, R. Soc. Sci. Upsal. 
Actis III. 1: 108. 1851; Sacc. Syll. Fung. 6:533. 1888; 
Lloyd, Myc. Writ. 4. Stip. Stereums, 34. 1913. 



3 



[Vol. 7 
114 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Type : a fragment in Kew Herb., according to Lloyd. 

Erect, cespitose, membranaceous-soft, fragile when dry, 
palmately branched, complanate, ribbed, dilated above, lacer- 
ate-fimbriate at the apex; hymenium definitely inferior, pallid 

gilvus; spores white. 
In pine woods, Pico de Orizaba, 10,000 ft. altitude, Mexico. 

Collected by Liebman. 

An extraordinary species, similar to Thelephora tuberosa and 
Tremellodendron pallidum but with the substance thin, some- 
what membranaceous, fragile when dry, and with the pileus 
foliaceous-complanate, ribbed (ribs commonly simple as in 
Alaria), very distinct. More than an inch high. Hymenium 
occupying the whole lower surface, at length floccose-collapsing 
and often foveolate, almost porose; basidia evidently 4-spored. 

The above is a translation of the original description. I did 
not find the type in Herb. Fries at Upsala nor see the frag- 
ment which Lloyd has reported as preserved at Kew. 

The specimen from Dutch Guiana, which is cited below, is so 
similar in aspect to Tremellodendron pallidum that it is probably 
S. craspedium. This cluster is 7 cm. in diameter and 3-4 cm. 
high, and agrees well with details of the original description. 
The basidia are simple, only detached spores found. These are 
hyaline, even, globose, 3 n in diameter. 

Specimens examined: 
Dutch Guiana: Jacob Samuels (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56300). 



26. S. petalodes Berkeley, Ann. & Mag. Nat. Hist. II. 9: 198. 

1852; Sacc. Syll. Fung. 6: 557. 1888; Massee, Linn. Soc. 

Bot. Jour. 27:165. 1890; Lloyd, Myc. Writ. 4. Stip. Stere- 
ums, 32. text f. 551. 1913. Plate 3, fig. 23. 

Illustrations: Lloyd, loc. cit. 

Type: in Kew Herb, according to Lloyd. 

Pileus coriaceous, sessile, at first infundibuliform, soon split 
into numerous lobes which are again more or less divided, dull 
reddish brown, marked with long grooves or striae; hymenium 
pale, much cracked, sometimes so much so as to be nearly 
granulated. 

San Domingo. Coll., Salle, 52. 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



115 



The above is the original description of 8. petalodes, a species 
of which I have seen no specimen. Lloyd's figure of the type 
shows the fructification to be a rosette-shaped mass 4 cm. high 
and 6 mm. in diameter, composed of many elongated pileate 
flaps, each of which is flattened and up to 7 mm. broad. No 
record was published by Berkeley as to whether S. petalodes 
grows on ground or on wood. 



27. S. anastomosans (Berk. & Curtis) Lloyd, Myc. Writ. 



4. Stip. Stereums, 35. 



1913. 



Thelephora anastomosans Berkeley & Curtis, Linn. Soc. Bot. 

Jour. 10: 329. 1868; Sacc. Syll. Fung. 6:534. 1888. 

Type: in Curtis Herb, and Kew Herb. 

Fructification stipitate, white, with the pileus divided into 
many segments; pileate branches and branchlets more or less 



laterally grown together above, somewhat flabel- 
liform and fimbriate, below more or less distinct 
or confluent into the common stem; hymenium 
even, inferior; no cystidia nor gloeocystidia ; 
spores copious, hyaline, even, subglobose, 4-4^- X 







O 



o 



3^-4 



2 



A* 



Fructifications about 2\ cm. high. 

On stump. Cuba. October. 

It was noted by the authors of the species that 



Fig. 8. 

S. anastomosans. 

Spores X 665. 

From type. 



S. anastomosans is allied to S. craspedium, but the divisions of 
its pileus are narrower than I understand them to be in the lat- 
ter. S. anastomosans is somewhat suggestive of S. Hartmanni 
and S. proliferum but differs in having many pileate divisions 
grow out from a common trunk so as to form a rosette-like 
mass, as in doubled forms of Thelephora caryophyllea. 

Specimens examined: 
Cuba: C. Wright, 280, type (in Curtis Herb.). 



28. S. proliferum (Berk.) Lloyd, Myc. Writ. 4. Stip. Stere- 



ums, 34. text f. 554. 1913. 



Plate 4, fig. 24. 



Thelephora prolifera Berkeley, Hooker's Jour. Bot. 8:272. 
1856; Sacc. Syll. Fung. 6: 542. 1888. 
Illustrations: Lloyd, loc. cit. 
Type: in Kew Herb, and Curtis Herb. 



• 



[Vol. 7 
116 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Fructifications cespitose, stipitate, coriaceous, erect, white, 
now between light buff and cartridge-buff throughout; stem 
cylindric, branched above, the branches either slender, cylindric, 
sterile bodies, or flattened, membranous pilei 1-2 mm. broad, 
1-1 1 cm. long; hymenium on the lower side, even; a few de- 
tached spores hyaline, even, 3|X3 m, none found on basidia. 

Fructifications about 3 cm. high; stems \ mm. in diam.; 
pileate branches 1-1 \ cm. long, 1-2 mm. broad. 

On roots of trees. Brazil. 

Berkeley described S. proliferum as somewhat creeping and 
having the branches with tips attached again to the matrix by 
means of large, orbicular, radiated and laciniated disks. These 
characters should render this species easy for the collector to 
recognize, but the herbarium specimen which I studied did not 
show the above feature noticeably; it had somewhat the aspect 
of S. Hartmanni but without the pectinate margins of the latter. 
The hymenium of the specimen studied is in poor condition and 
the spore characters, as given above, are uncertain. I studied 
for N. Y. Bot. Gard. Herb., No. 508, a fungus collected at 
Church Cove, Bermuda, which has the general aspect of S. 
proliferum but with spores hyaline, even, 13-16X6-7 n, and is 
probably a distinct species. Still it is well to keep S. proliferum 
in mind in connection with species of the West Indies. 

Specimens examined: 
Brazil: Rio Negro, Spruce, 17, type (in Curtis Herb.). 



29. S. caespitosum Burt, n. sp. Plate 4, fig. 25. 

Type: in Burt Herb. 

Fructifications coriaceous, thin, cespitose, effuso-reflexed, 
with the resupinate portion small and bearing a cluster of broader 
and longer, imbricate, pileate lobes which are somewhat fur- 
furaceous or with minute t omentum on the upper side, glabrate 
towards the margin, drying tawny and zonate with ochraceous, 
tawny zones, the margin entire; hymenium even, whitish to 



buff 



intermediate 



layer bordered above by a narrow, slightly colored zone and 
composed of densely longitudinally arranged, hyaline, thick- 
walled hyphae 3^ /j. in diameter; hymenial layer up to 120 n 
thick, containing numerous slender, flexuous gloeocystidia 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



117 



Q 




3 §-5 n in diameter near the base, tapering out- 
ward; no colored conducting organs nor note- 
worthy paraphyses; spores hyaline, even, 4-4|X 
3-3| fi, copious. 

Resupinate portion covers area 6X5 mm., re- 
flexed lobes 5-10 mm. in diameter — about 10 in 
the cluster. 

On broken lateral stub of dead limb of a fron- 
dose species. Jamaica. January. Probably rare. 

Viewed from above, S. caespitosum has the gen- 
eral aspect and coloration of species of Stereum 
in sections having stems, as S. pergamenum and 
S. decolorans, but is excluded from these sections 
by attachment to the substratum by a distinctly 
resupinate portion. The species is unique in the 
effuso-reflexed section in the above resemblance, 
and with additional characters of clustered, 
imbricated habit of growth and presence of 
gloeocystidia, should be readily recognized. 

Specimens examined: 
Jamaica: Moneague to Union Hill, W. A. Murrill, 1181, type, 



Fig. 9. 

S. caespitosum. 
Gloeocystidia 

and spores X 

665. 
From type. 



comm. by N. Y. Bot. Gard. Herb. 

. S. fuscum Schrader ex Quelet, Fl. Myc. France 



Bresadola, I. R. Accad. Aedati Atti III 



1888; 



Thelephora f 



1897. 

Plate 4, fig. 26 



Syn 



Schrader, Spic. Fl. Germ. 184. 1794; 



both places renaming: the 



and Myc 



My 

Syn 
um 



um 



Eur. i: 122. 1822 
T. bicolor); Fries, Syst. 



i:438. 1821 (following Persoon).— T. bicolor Persoon, 



Fung. 568. 



Syst. My 



Thelephora); Fries, Epicr. 549. 
Morgan, Cincinnati Soc. Nat. Hist 



58. 1821.— Store 

(under **** Stere 



1838; Hym. Eur 



Sacc 



Syll 



Fung. 6:565. 1888; Massee, Linn. Soc 



1888; 
Bot. 



Jour. 27: 177. 1890— £. coffeatum Berk. & Curtis, Grevillea 

: 568. 1888; Massee, Linn. 



1: 164. 1873; Sacc. Syll. Fung. 6 
Soc. Bot. Jour. 27: 190. 

Illustrations: Fries, Icones Hym. pi. 197. f. 2: Karsten 



1890. 



Hym 



/ 



118 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Fructifications somewhat membranaceous 



soft 



spongy 



sometimes resupinate, usually becoming conchate-reflexed, ofter 

imbricated, villose, becoming glabrous 




somewhat 



sulcate, dry 



glab 



brown to bister: hymenium even 



white, dry 



cream 



to 



pallid mouse 



in structure 1000 n 
longitudinally and 



3 



a diam- 
surface, 



thick, composed of lonj 
loosely interwoven hypha 
eter, colored towards the 
hyaline towards the hymenium; hyme- 
nium not zonate, containing flexuous 
gloeocystidia 20-60X5-7 n, rarely 90 n, 



hyaline, 3 



3 



j" 



Reflexed Dileus 1-4 cm. lone, 2-5 cm 



cm 



Fig. 10. S. fuscum. 
Gloeocystidia and spores 
X 665. 



April to December 



wide ; resupinate specimens 3-10 X 1 

On rotting frondose limbs usually, but 
sometimes on pine. Canada to Texas, 
westward to Oregon, in the West Indies, 
and also in Europe. 
Not rare. 
Reflexed specimens of S. fuscum may be recognized at sight 
by the soft, pliant pileus, brown and felt-like above, with a white 
hymenium. Gloeocystidia are so rare in the hymenium of a 
Stereum that their presence in abundance in this species affords 
a decisive specific character. Wholly resupinate specimens 
have the color of the hymenium of reflexed fructifications and 
have similar consistency and gloeocystidia. So many reflexed 
species occur resupinate that one should be sure to gather the 
more or less reflexed fructifications which can usually be secured 
associated with the resupinate specimens. Since both Persoon 
and Fries recognized the priority of Schrader's specific name fus- 
cum and substituted bicolor, presumably because highly distinc- 
tive and appropriate for the species, the restoration of the 
original name by recent mycologists seems just. 

Specimens examined: 
Exsiccati: Ellis, N. Am. Fungi, 1207; Ell. & Ev., Fungi Col., 

1019; Rabenhorst, Fungi Eur., 3233; Ravenel, Fungi Am., 
9; Ravenel, Fungi Car. 2:33; de Thiimen, Myc. Univ., 
1704. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 119 

Finland: Mustiala, P. A. Karsten, in de Thumen, Myc. Univ., 

1704. 
Sweden: Femsjo, L. Romell, J+02. 
England: Selby, E. A. Burt. 
France: Allier, H. Bourdot, 1611+1. 
Hungary: Kmet, comm. by G. Bresadola. 
Canada: J. Macoun, 76, 280. 
Ontario: Ottawa, J. Macoun, 21, 59; Toronto, J. H. Fault, 

Univ. Toronto Herb., 361 (in Mo. Bot. Gard. Herb., 

44863). 
Vermont : Middlebury, E. A . Burt; North Ferrisburg, E. A . Burt. 

New York: Bronx Park, New York, H. D. House (in N. Y. 

State Mus. Herb, and Mo. Bot. Gard. Herb., 54392), 

and W. A. Murrill (in N. Y. Bot. Gard. Herb, and Mo. 

Bot. Gard. Herb., 56773); Staten Island, W. H. Ballou 

(in Burt Herb., N. Y. Bot. Gard. Herb., and Mo. Bot. 

Gard. Herb., 56774) ; Syracuse, D. C. Mills (in N. Y. Bot. 

Gard. Herb, and Mo. Bot. Gard. Herb., 56281). 
Pennsylvania: Kittanning, D. R. Sumstine; West Chester, Ever- 

hart & Haines, in Ellis, N. Am. Fungi, 1207. 
District of Columbia: C. L. Shear, 1039; Takoma Park, C. L. 

Shear, 951+. 

South Carolina: H. W. Ravenel, in Ravenel, Fungi Car. 2: 33; 

Santee Canal, H. W. Ravenel, 910 (in Curtis Herb.), and 
Curtis Herb., 2923, type of Stereum coffeatum (in Kew 
Herb.); Salem, Schweinitz (in Herb. Schweinitz). 

Georgia: Atlanta, E. Bartholomew, 5680 (in Mo. Bot. Gard. 

Herb., 44219); Tipton, C. J. Humphrey, 156. 
Florida: Gainesville, H. W. Ravenel, in Ravenel, Fungi Am., 

9; Lake City, P. L. Richer, 898; New Smyrna, C. G. 

Lloyd, 2118. 

Alabama: Auburn, L. M. Underwood, comm. by U. S. Dept. 

Agr. Herb., F. S. Earle (in Mo. Bot. Gard. Herb., 5058), 
and F. S. Earle & C. F. Baker; Fayette Co., P. V. Siggers, 
comm. by A. H. W. Povah, 15 (in Mo. Bot. Gard. Herb., 
9226); Montgomery Co., R. P. Burke, S3 (in Mo. Bot. 
Gard. Herb., 15763). 

Mississippi: Chicou (in Mo. Bot. Gard. Herb., 43014). 

Louisiana: Abita Springs, A. B. Langlois; New Orleans, F. S. 






i on ' VoL - 7 

1/U ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Earle (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 
56775); St. Martinville, A. B. Langlois, bz, 2095, and a 
specimen comm. by Lloyd Herb., 2737. 

Texas: San Antonio, W. H. Long, 21703 (in Mo. Bot. Gard. 

Herb., 55164). 

Ohio: A. P. Morgan (in Lloyd Herb.) and C. G. Lloyd, in Ell. 

& Ev., Fungi Col., 1019; Linwood, C. G. Lloyd, 1154, 1326; 
Norwood, C. G. Lloyd, V. 

Indiana: Greencastle, L. M. Underwood (in N. Y. Bot. Gard. 

Herb, and Mo. Bot. Gard. Herb., 56276, 56278); Hibernian 

Mills, Whetzel & Reddick, comm. by D. Reddick, 3. 
Wisconsin: Madeline Island, V. B. Walker, 6a (in Mo. Bot. 

Gard. Herb., 8359); Madison, Miss A. 0. Stucki, 26. 
Missouri: Marianna, H. von Schrenk (in Burt Herb, and Mo. 

Bot. Gard. Herb., 42836); Oran, H. von Schrenk (in Mo. 

Bot. Gard. Herb., 42835); Perryville, C. H. Demetrio, in 

Rabenhorst, Fungi Eur., 3233; Williamsville, B. M. Duggar, 
482. 

Arkansas: Cass, W. H. Long, 19923 (in Mo. Bot. Gard. Herb., 

13266); Levisque, P. Spaulding (in Mo. Bot. Gard. Herb., 

5057). 

Idaho: Kooskia, J. R. Weir, 589 (in Mo. Bot. Gard. Herb., 

56776). 

British Columbia: Agassiz, /. R. Weir, 603 (in Mo. Bot. Gard. 

Herb., 36748). 

Oregon: Corvallis, C. E. Owens, 2037 (in Mo. Bot. Gard. Herb., 

43871). 

Cuba: Alto Cedro, L. M. Underwood & F. S. Earle, 1571, 1581, 

comm. by N. Y. Bot. Gard. Herb.; Baracoa, L. M. Under- 
wood & F. S. Earle, 504, comm. by N. Y. Bot. Gard. Herb. 

Jamaica: Cinchona, W. A. & E. L. Murrill, 462, comm. by N. Y. 

Bot. Gard. Herb. ; Hope Gardens, F. S. Earle, 500, comm. 
by N. Y. Bot. Gard. Herb.; Mandeville, A. E. Wight, 
comm. by W. G. Farlow; Troy and Tyre, W. A. Murrill 
& W. Harris, 1073, comm. by N. Y. Bot. Gard. Herb. 



31. S. rufum Fries, Epicr. 553. 1838; Hym. Eur. 644. 1874; 
Sacc. Syll. Fung. 6:575. 1888; Romell, Bot. Not. 1895: 



71. 1895 



fig 






1920] 



HURT — THELEPHORACEAE OF NORTH AMERICA. XII 



121 



Thelephora rufum Fries, Elenchus Fung, i : 187. 1828. 
Cryptochaete rufa (Fries) Karsten, Finska Vet.-Soc. Bidrag 
Natur och Folk 48: 408. 1889.— Tuber cularia pezizoidea 
Schweinitz. 



Syll. Fung. 



Am. Phil. Soc. Trans. N. S. 4:301. 1832; Sacc 



644 



1886. 



Hypocrea Richardsonii Berkeley 



& Montaene, Grevillea 



1875; Sacc. Syll 



1883; Ellis & Everhart, N. Am. Pyrenom 



86. 



1892. 



Corticium pezizoideum 
Bui. 21: 385. pi. 218. 



(Schw 
1894. 



Schrenk, Torr. Bot. Club 



Illustrations: von Schrenk, Torr. Bot. Club 



1894 



Type: in Herb. Fries. 

Fructifications scattered or gregarious, coriaceous-fleshy, 
bursting out from the bark, verruciform, plicate-tuberculose, 










<*** 







S0 









Fig. 11. S. rufum. Fructifications, /; section of fructification, m; section of 
hymenial region, n; spores, s. After von Schrenk. 



peltate, vinaceous-brown to hematite-red, under side glabrous, 
the margin free all around; hymenium becoming coarsely 
wrinkled, vinaceous-brown, often grayish pruinose; in structure 
2 mm. thick at the center, 600-800 n thick in the marginal 



1- 

portion, composed of ascending, loosely interwoven, incrusted, 
hyaline hyphae 4-4 \ n in diameter over the incrustation; flexuous 
gloeocystidia 50-90X7-10 p. are scattered in or near the hymeni- 
um but not protruding; spores white in spore collection, even, 
curved, 6-8X1^-2 p. 

Fructifications 2-4 mm. in diameter. 

On dead fallen Populus tremuloides. Newfoundland to Mass- 
achusetts and westward to North Dakota and Colorado. 
March to December. Common. Occurs in Scandinavia also. 



[Vol. 7 



122 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



aS. rufum may be recognized at sight by its occurrence on 
prostrate poplar limbs and logs in the form of small vinaceous 
fructifications with the hymenium gyrosely wrinkled. The 
fructifications become peltate when full grown, attached by the 
center, and with the marginal portions free and turned outward. 

Specimens examined: 
Exsiccati: Bartholomew, Fungi Col., 1817, under the name 

Corticium rufo-marginatum, and 2716; Ellis, N. Am. Fungi, 
1329; Romell, Fungi Scand. Exs., 123; Shear, N. Y. Fungi, 

88. 
Norway: Christiania, M. N. Blytt, authentic specimen (in 

Herb. Fries). 
Sweden: Stockholm, L. Romell, in Romell, Fungi Scand. Exs., 

123; Upsala, L. Romell, 39. 
Newfoundland: B. L. Robinson & H. von Schrenk (in Mo. Bot. 

Gard. Herb., 42944); Bay of Islands, A. C. Waghorne (in 

Mo. Bot. Gard. Herb., 17692). 
Ontario: Toronto, T. Langton, Univ. Toronto Herb., 595 (in 

Mo. Bot. Gard. Herb.). 
Maine: Orono, F. L. Harvey, 6 (in Mo. Bot. Gard. Herb., 

16620); Portage, L. W. Riddle, 10. 
New Hampshire: Shelburne, W. G. Farlow (in Mo. Bot. Gard. 

Herb., 14796). 
Vermont: Middlebury, E. A. Burt, two collections; North Fer- 

risburg, E. A. Burt. 

Massachusetts: Peabody, A. R. Sweetser; Waverley, H. von 

Schrenk (in Mo. Bot. Gard. Herb., 16623). 

New York: Alcove, C. L. Shear, in Shear, N. Y. Fungi, 88; 

East Galway, E. A. Burt; Ithaca, G. F. Atkinson (in Mo. 
Bot. Gard. Herb., 4775); Willsboro Point, C. 0. Smith, in 
Bartholomew, Fungi Col., 1817. 

Pennsylvania: Trexlertown, W. Herbst. 

Michigan: Mackinac Island, E. T. & S. A. Harper, 707; North- 
port, H. von Schrenk (in Mo. Bot. Gard. Herb., 22481). 

Wisconsin: La Crosse, W. Trelease (in Mo. Bot. Gard. Herb., 

14794); Madison, W. Trelease, in Ellis, N. Am. Fungi, 
1329, and (in Mo. Bot. Gard. Herb., 14794, 16621); Pal- 
myra, Miss A. 0. Stucki, 27; Syene, W. Trelease, 3022 (in 
Mo. Bot. Gard. Herb., 14793). 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



123 



Nebraska: Lincoln, Miss L. B. Walker, 7 (in Mo. Bot. Gard. 

Herb., 44818). 
North Dakota: Fargo, F. J. Seaver, 25, 54 (in Mo. Bot. Gard. 

Herb., 16222, 16637). 
Montana: Helena, F. W. Anderson, 202 (in Mo. Bot. Gard. 

Herb., 21165). 
Colorado: Blind Canon Placer, C. L. Shear, 1021; Golden, E. 

Bartholomew & E. Bethel, in Bartholomew, Fungi Col., 

2716, and E. Bethel & L. 0. Overholts, comm. by L. 0. 

Overholts, 1754 (in Mo. Bot. Gard. Herb., 54875) ; Ouray, 

C. L. Shear, 1187. 

32. S. Pini Fries, Epicr. 553. 1838; Hym. Eur. 643. 



Sacc. Syll. Fung. 6: 574. 1888. 



1874; 
Plate 4, fig. 28. 

Thelephora Pini Fries, Syst. Myc. 1: 443. 1821; Elenchus 
Fung. 1 : 187. 1828. — Sterellum Pini (Schleich.) Karsten, Finska 
Vet.-Soc. Bidrag Natur och Folk 48: 405. 1889. 

Illustrations: Smith, Brit. Basidiomycetes, text/. 98 E, F. 

Fructifications gregarious, coriaceous-cartilaginous, orbic- 
ular, resupinate, with the margin free and attached by the 
center, shield-shaped, finally bullate, drying rigid, Benzo-brown; 
hymenium wood-brown to Benzo-brown, somewhat pruinose, 
becoming somewhat 
tuberculose; in struc- 
ture 500 fji thick, thin- 
ning out towards the 
margin, with the in- 
termediate layer bor- 
dered on each side by 
a narrow, colored zone , 
and composed of lon- 
gitudinally arranged, 

y , ' Fig. 12. S. Pini. Fructifications, /, natural size; 

hyaline hyphae with cystidia, c, and gloeocystidia, g, X 665. 

walls gelatinously 

modified, the subhymenium olivaceous-colored; cystidia in- 
crusted, 24X8 n, sometimes very few to be found; fusoid or 
irregular gloeocystidia, 30-40X10-15 m> are sparingly present 
in or near the hymenium; spores hyaline, even, curved, 5-6 
X2-2£ M . 







[Vol. 7 
124 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Fructifications 1-4 mm. in diameter. 

On bark of fallen limbs of Pinus resinosa. Maine and New- 
Hampshire. August. Rare. 

The fructifications are so near the color of the bark of the 
dead pine limbs upon which they grow that they are likely to be 
overlooked, or, if collected, roughly classed among the Dis- 
comycetes on account of their resemblance to these fungi in 
aspect. The occurrence on pine bark, small, shield-shaped 
fructifications Benzo-brown in color, and showing in section 
both cystidia and gloeocystidia are a combination of characters 
which should not fail to identify this species. 

Specimens examined: 

Exsiccati: Krieger, Fungi Sax., 364; Rabenhorst, Herb. Myc, 

213. 

Finland: Mustiala, P. A. Karsten. 
Sweden: Stockholm, L. Romell, 82. 
Germany: Dresden, in Rabenhorst, Herb. Myc, 213; Konigs- 

tein, Saxony, W. Krieger, in Krieger, Fungi Sax., 364. 
France: St. Priest, Allier, H. Bourdot, 15067. 
Maine: /. Blake, 659 (in Curtis Herb.). 
New Hampshire : Chocorua, W. G. Farlow, 37. 



33. S. purpureum Persoon, Roemer Neues Mag. Bot. 1: 
110. 1794; Obs. Myc. 2:92. 1799; Fries, Epicr. 548. 1838; 
Hym. Eur. 639. 1874; Berkeley, Brit. Fung. 270. 1860; Mor- 
gan, Cincinnati Soc. Nat. Hist. Jour. 10: 194. 1888; Sacc. 
Syll. Fung. 6: 563. 1888; Massee, Linn. Soc. Bot. Jour. 27: 
186. 1890. Plate 4, fig. 29. 

Thelephora purpurea Persoon, Syn. Fung. 571. 1801; Myc. 
Eur. 1: 121. 1822; Fries, Syst. Myc. 1: 440. 1821.— Stereum 
vorticosum Fries, Obs. Myc. 2: 275. 1818; Epicr. 548. 1838; 
Hym. Eur. 639. 1874; Sacc. Syll. Fung. 6: 563. 1888. 

Illustrations: Fl. Danica3: pi. 534-f. 4', Hussey, 111. Br. Myc. 
pi. 20. f. A; Istvanffi, Jahrbuch. f. wiss. Bot. 29: pi. 6.f. 37-39; 
Lanzi, Fungi di Roma, pi. 11. f. 2: Sowerby, Col. Figs. Eng. 
Fungi, pi. 388. f. 1. 

Type: authentic specimen from Persoon in Kew Herb. 
Fructifications coriaceous-soft, drying rigid, sometimes resup- 
inate, usually more or less reflexed, often imbricated, the 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 125 

upper side villose-tomentose, light buff to cartridge-buff, the 
margin entire; hymenium even, glabrous, light purple-drab to 
dark vinaceous-drab ; in structure about 500-800 n thick ex- 
cluding the tomentum, with the intermediate layer more looselv 



ed on its under side 



subhymeni 



orm, orsubelob 




T 









ular organs 15-30X12-25 M ; no W^/fS^ 
cystidia; spores hyaline, even, ^sz'c&fkP 
flattened on one side, 5-7 X 2§-3 /*. 
Fructifications with resupinate 
portion about 1-2 cm. in diam- 
eter; reflexed portion 5-20 mm. Fi &- 1:J - S - purpureum. Section 
broad and sometimes orisneH of h^rml region X 90, and vesic- 

oroaa, ana sometimes cnspea ukr bod[es x 665 From authentic 

or lobed with lobes 5 mm. in specimen. 

diameter. 

On dead stumps and logs of Populus, Betula, and other fron- 
dose species. Newfoundland to Delaware and westward to 
British Columbia and Oregon, also in Uruguay and in Europe. 
June to April. Common but not ranging into torrid regions. 

S. purpureum is usually recognized by its buff, tomentose 
pileus, purplish hymenium which does not bleed when wounded. 



d 



Sectional preparations show charac 



teristic vesicular organs in the subhymenial region, such as 
are present in the closely related S. rugosiusculum, but no hair- 
like cystidia in the hymenium, by the absence of which S. pur- 
pureum is distinguished from the latter. 

The authentic specimen of S. vorticosum in Herb. Fries at 
Upsala is 2-3 X If cm., narrowly reflexed, with dark purplish 
hymenium, and with the usual microscopic structure and spores 
of S. purpureum. 

Specimens examined: 
Exsiccati: Bartholomew, Fungi Col., 3489; Berkeley, Brit. 

Fungi, 147; Cooke, Fungi Brit., 12; Ell. & Ev., N. Am. 
Fungi, 2018, 2601; Klotzsch, Fungi Germ., 50; Krieger, 
Fungi Sax., 1852; Rabenhorst, Herb. Myc, 504; Romell, 
Fungi Scand. Exs., 27; Shear, N. Y. Fungi, 311. 
Europe: authentic specimen of Thelephora purpurea from Per- 

soon (in Herb. Hooker in Kew Herb.). 



[Vol. 7 
126 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Sweden: E. Fries (in Kew Herb.); Femsjo, authentic specimen 

of Stereum vorticosum (in Herb. Fries) ; Stockholm, L. Ro- 
mell, 34, 288, and in Romell, Fungi Scand. Exs., 27. 

England: M. J. Berkeley, in Berkeley, Brit. Fungi, 147; Hamp- 

stead, M. C. Cooke, in Cooke, Fungi Brit., 12. 

France: Corrombles, comm. by Lloyd Herb., 3355; St. Priest, 

Allier, H. Bourdot, 12459, 12461. 

Germany: Klotzsch, in Klotzsch, Fungi Germ., 50; Dresden, 

in Rabenhorst, Herb. Myc, 504; Winterberge, Wagner & 
Krieger, in Krieger, Fungi Sax., 1852. 

Austria: Stapf, Fl. Exs. Austro-Hungarica, 3543 (in Mo. Bot. 

Gard. Herb., 5125, 715171). 

Italy: Trento, G. Bresadola. 

Newfoundland: Bay of Islands, A. C. Waghorne, 20, 86 (in Mo. 

Bot. Gard. Herb., 5091, 5092). 

Ontario: Harraby, E. T. & S. A. Harper, 64U Ottawa, J. 

Macoun, 17, 39; J. M. Macoun, comm. by N. Y. State 
Mus. Herb, (in Mo. Bot. Gard. Herb., 56085); Port 
Credit, J. H. Fault, Univ. Toronto Herb., 646 (in Mo. Bot. 
Gard. Herb., 44944); Toronto, R. P. Wodehouse, J. H. 
Fault, G. H. Graham, Univ. Toronto Herb., 310, 311, 677, 
respectively (in Mo. Bot. Gard. Herb., 44887, 44889, 
44920); Wilcox Lake, /. H. Fault, Univ. Toronto Herb., 
377 (in Mo. Bot. Gard. Herb., 44929). 

Maine: Manchester, F. L. Scribner, comm. by U. S. Dept. Agr. 

Herb.; Orono, F. L. Harvey, 3 (in Mo. Bot. Gard. Herb., 
43850) and in Ell. & Ev., N. Am. Fungi, 2018; Portage, 
L. W. Riddle, 6. 

Vermont: Brattleboro, E. A. Burt; Little Notch, E. A. Burt; 

Middlebury, E. A. Burt, three collections; North Ferris- 
burg, E. A. Burt; Ripton, E. A. Burt, three collections; 
Walden, L. S. Orton, 4 (in Mo. Bot. Gard. Herb., 44081). 

Massachusetts: Cambridge (in Mo. Bot. Gard. Herb., 5094). 



Connecticut: C. C. Hanmer, 2326, 2061 (in Mo. Bot. Gard. 

Herb., 43847/8). 

New York: Sartwell (in Mo. Bot. Gard. Herb., 5151, 5156); 

Alcove, C. L. Shear, 1120, 1122, and in Shear, N. Y. Fungi, 
311; East Galway, E. A. Burt; Ithaca, G. F. Atkinson, 
2093, 2141, C. J. Humphrey, 307, H. S. Jackson & C. Lewis, 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 127 



19396; Long Lake, A. H. W. Povah (in Mo. Bot. Gard. 
Herb., 9227); North Elba, C. H. Kaufman, 8 (in Mo. Bot. 
Gard. Herb., 16701); Rome, H. von Schrenk (in Mo. Bot. 
• Gard. Herb., 55022, 55024/5). 

Pennsylvania: Bethlehem, Schweinitz (in Herb. Schweinitz); 

Trexlertown, W. Herbst, 16, 28, and coram, by Lloyd Herb., 
3603. 

Delaware: Wilmington, A. Commons, in Ell. & Ev., N. Am. 

Fungi, 2601. 

Ohio: Norwood, C. G. Lloyd, 1787, and (in Mo. Bot. Gard. Herb., 

5093). 
Indiana: Indianapolis, J. B. Demaree, comm. by G. W. Hoffer 

(in Mo. Bot. Gard. Herb., 54790); Lafayette, C. R. Orton, 

5 (in Mo. Bot. Gard. Herb., 44082). 
Wisconsin: Madison, W. Trelease (in Mo. Bot. Gard. Herb., 

5043); Star Lake, Miss A. O. Stucki, Univ. Wis. Herb., 59. 
Minnesota: Park Rapids, comm. by E. L. Jensen, 10 (in Mo. 

Bot. Gard. Herb., 11100). 
Montana: Helena, Monarch, J. R. Weir, 587, 598 (in Mo. Bot. 

Gard. Herb., 56738, 58739). 
Wyoming: Boulder, F. S. Wolpert, comm. by J. R. Weir, 7949 

(in Mo. Bot. Gard. Herb., 56219). 
Idaho: Priest River, /. R. Weir, 10. 
British Columbia: Sidney, /. Macoun, 74- (in Mo. Bot. Gard. 

Herb., 55352); Vancouver Island, /. Macoun, 51 (in Mo. 

Bot. Gard. Herb., 5737), and comm. by J. Demaree, V88 

(in Mo. Bot. Gard. Herb., 22752). 
Washington: Bingen, W. N. Suksdorf, 766. 767; Easton, C. J. 



Humphrey, 644-9; Olympia, C. J. Humphrey, 6292; Seattle, 

S. M. Zeller, 108 (in Mo. Bot. Gard. Herb., 44140). 
Oregon: Corvallis, C. E. Owens, 2076 (in Mo. Bot. Gard. Herb., 

44038). 
Uruguay: Montevideo, W. Mitten Herb., 1325 (in N. Y. Bot. 

Gard. Herb, and Mo. Bot. Gard. Herb., 56691). 



34. S. rugosiusculum Berk. & Curtis, Grevillea 1: 162. 1873; 
Morgan, Cincinnati Soc. Nat. Hist. Jour. 10: 193. 1888; Sacc. 
Syll. Fung. 6: 567. 1888; Massee, Linn. Soc- Bot. Jour. 27: 
187. 1890. Plate 4, fig. 30. 



128 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Stereum Micheneri Berk. & Curtis emend. Massee, Linn. Soc. 
Bot. Jour. 27: 183. 1890. — S. Micheneri Berk. & Curtis, Gre- 
villea 1: 162. 1873 (in part). See Ann. Mo. Bot. Gard. 1: 



214. 



1914. — Corticium Nyssae Berk. & Curtis, Grevillea 1 : 166. 
1873; Sacc. Syll. Fung. 6: 609. 1888; Massee, Linn. Soc. Bot. 



Jour. 27: 120. 



1890. 



C. siparium Berk. & Curtis, Grevillea 




^ s 




m, 




1: 177. 1873; Sacc. Syll. Fung. 6:636. 1888; Massee, Linn. 

Soc. Bot. Jour. 27: 139. 1890. 

Illustrations: Berkeley, Ann. & Mag. Nat. Hist. I. 1: 94. 

pi. 5. f. J+5. 

Type: in Kew Herb, and Curtis Herb. 

Fructifications coriaceous-soft, rarely resupinate, usually more 

or less broadly reflexed, upper surface tomentose, spongy, some- 
times with projecting hairs 
collapsed together into a 
plane or wrinkled surface, 
drying cartridge-buff to 
cinnamon-buff, the margin 
entire; hymenium even, 
drying vinaceous-buff to 
fawn color; in structure up 
to 1-1| mm. thick inclusive 
of the tomentum, with the 
intermediate layer on its 
under side in the sub- 

hymenial region, loosely 

interwoven, and containing more or less numerous, pyriform 
vesicular bodies 15-30X10-20 /*; cystidia slender, thin-walled, 
tapering hairs, not incrusted, 4-5 n in diameter, protruding up 
to 25 n beyond the basidia; spores white in spore collection, 
even, flattened on one side, 4§-6X2-3 m- 

Resupinate specimens up to 6 cm. in diameter; reflexed por- 
tion 1-2 cm. broad, 2-6 cm. laterally along substratum. 

On logs and stumps of Salix and other frondose species. 
Ontario to Alabama, in Missouri, and in British Columbia to 
Mexico; occurs also in Sweden, France, Italy, England, and 
Japan. August to April. 

Stereum rugosiusculum is probably more frequent and more 
widely distributed than shown by the specimens received, for 



Fig. 14. S. rugosiusculum. Section of hy- 
menial region X 90; cystidium and basidia, 
n, vesicular body, v , and spores, s, X 665. 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 129 



the general aspect and microscopic structure of specimens are 
usually so similar to S. purpureum that it is distinguishable from 
the latter only by the presence of weak flexuous hairs in the 
hymenium which are not visible until sectional preparations are 
examined with the compound microscope. Such hymenial hairs 



were in 1839 figured by Berkeley, loc. cit., in illustrating the 
hymenium of what he regarded as Thelephora purpurea but 
which now appears to have been S. rugosiusculum. All speci- 
mens in which these hair-like cystidia have been demonstrated 
have been either resupinate or with simple, re flexed portion 
not narrowly lobed or complicate. It has not been possible to 
observe a specimen throughout its whole season of growth to 
determine whether the hair-like cystidia are a constant character. 
In forming the glabrous, rugulose surface upon which the specific 
name is based, the specimens do not become denuded of their 
original tomentose covering, for sectional preparations of such 
specimens, mounted in liquid medium, show this hairy covering 
to be of the original thickness and with the tips of the hairs no 
longer adhering together into a plane surface but now floating 
free. Probably the gluing together of the hairs into a glabrous 
surface is a weather phenomenon. 

Specimens examined: 
Exsiccati: Bartholomew, Fungi Col., 3489, under the name Ster- 

eum purpureum; Cavara, Fungi Longobardiae, 60, under 
the name Stereum purpureum; Ellis, N. Am. Fungi, 323, 
under the name Stereum purpureum. 
Sweden: Stockholm, L. Romell, 83. 

England: M. J. Berkeley, under the name Stereum vorticosum 

(in Kew Herb.). 

France: Fautrey, determined by Patouillard as S. purpureum, 

comm. by Lloyd Herb., 4339, 4363. 
Italy: F. Cavara, in Cavara, Fungi Longobardiae, 60. 
Ontario: London, J. Dearness, in Bartholomew, Fungi Col., 

3489. 

Maine: Morse, comm. by Sprague (in Curtis Herb., 5413, type 

of Stereum Micheneri as emended by Massee); Harrison, 
J. Blake, comm. by P. L. Picker; Piscataquis Co., W. A. 
Murrill, 1860, 2158 (in N. Y. Bot. Gard. Herb, and Mo. 
Bot. Gard. Herb.. 56692. 56693 s !. 



4 



130 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



Vermont: Ripton, E. A. Burt. 

Massachusetts: Sprague, 492, type (in Kew Herb, and Curtis 

Herb., 5412) ; Cambridge, H. von Schrenk (in Mo. Bot. 
Gard. Herb., 4774), and A. B. Seymour, T 19 (in Mo. Bot. 
Gard. Herb., 43886). 

New York: Ithaca, G. F. Atkinson, K, 2818a; Lake Placid, 

W. A. & E. L. Murrill, 445 (in N. Y. Bot. Gard. Herb, and 
Mo. Bot. Gard. Herb., 56694); White Plains, L. M. Under- 
wood (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 
56268) . 

New Jersey: J. B. Ellis, in Ellis, N. Am. Fungi, 323. 

Pennsylvania: E. Michener, 509, type of Corticium Nyssae (in 

Curtis Herb., 3486); Ohiopyle, W. A. Murrill, 1043 (in 
N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 56695); 
Trexlertown, W. Herbst. 

Virginia: Blacksburg, W. A. Murrill, 351 (in N. Y. Bot. Gard. 

Herb, and Mo. Bot. Gard. Herb., 56710). 

Alabama: Peters, 858, type of Corticium siparium (in Curtis 

Herb., 5239); Montgomery Co., R. P. Burke (in N. Y. Bot. 
Gard. Herb, and Mo. Bot. Gard. Herb., 56792). 

Missouri: Creve Coeur Lake, E. A. Burt (in Mo. Bot. Gard. 

Herb., 13031). 
Idaho: Priest River, J. R. Weir, 595 (in Mo. Bot. Gard. Herb., 

36740). 
British Columbia: J. Macoun, 62 (in Mo. Bot. Gard. Herb., 

5740) . 
Washington: Bellingham, J. R. Weir, 604 (in Mo. Bot. Gard. 

Herb., 36741) ; Seattle, W. A. Murrill, 129, 139, 147, comm. 
by N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 55743, 
55732, 55728); W. A. Murrill, 136, comm. by N. Y. Bot. 
Gard. Herb, (in Mo. Bot. Gard. Herb., 55735), and S. M. 
Zeller, 129 (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. 

Herb., 129). 

Oregon: Corvallis, W. A. Murrill, 892a, comm. by N. Y. Bot. 

Gard. Herb, (in Mo. Bot. Gard. Herb., 55724); Kiger 
Island, S. M. Zeller, 1788 (in Mo. Bot. Gard. Herb., 56653). 

California: R. A. Harper, 36 (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56697); Sierra Nevada Mts., Hark- 
ness, 1060 (in Herb. Cooke in Kew Herb., under the name 
Stereum muscigenum) . 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



131 



Mexico: Guernavaca, W. A. & E. L. Murrill, 410, 546, 547 (in 

N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 54535, 
54581, 54582). 

Japan: Kushiro, A. Yasuda, 64 (in Mo. Bot. Gard. Herb., 

56136). 



35. S. Murrayi (Berk. & Curtis) Burt, n. comb. 

Plate 4, figs. 31, 32. 
Thelephora Murraii Berk. & Curtis, Linn. Soc. Bot. Jour. 



10: 329. 1868; Grevillea 1 



150. 



1873; spelling of specific 

1888.— 



name changed to Murrayi in Sacc. Syll. Fung. 6: 546. 
Stereum tuberculosum Fries, Hym. Eur. 644. 1874; Sacc. Syll. 
Fung. 6: 586. 1888; Massee, Linn. Soc. Bot. Jour. 27: 204. 
1890; Romell, Bot. Not. 1895: 70. 1895. — S. pulverulentum 
Peck, Torr. Bot. Club Bui. 27: 20. 1900; Sacc. Syll. Fung. 16: 



187. 



1902. 



Illustrations: Lloyd, Myc. Writ. 5. Myc. Notes 62: pi 148. 



f. 1690. 



1920. 



Type: in Kew Herb, and Curtis Herb. 

Fructifications corky, adnate, usually resupinate and broadly 
effused, sometimes reflexed, the reflexed upper surface a hard, 
horny crust, not shining, concentric- 
ally sulcate, fuscous-black or aniline- 
black, the margin entire; hymenium 
drying from pale olive-buff to avella- 
neous, tubercular, deeply cracking; in 

structure 300 11 thick at first, then be- 
coming stratose and thickening to 800 
-2000 n, composed of densely inter- 
woven, rather suberect hyaline hyphae 
2^-4 n in diameter and of very numer- 
ous, hyaline, pyriform vesicular organs «,-■„„ . „ 

ik onOio iK u- V A' * -u\ A Flgl5> S. Murrayi. Sec- 

15-20X12-15 m which are distributed tion of hymenial region x 

throughout the whole fructification; no 488, showing vesicular bodies. 

colored conducting organs nor cystidia; 

spores white in spore collection, even, flattened on one side, 

4-5X2* m. 

Resupinate specimens 1-10 cm. in diameter, becoming con- 
fluent, reflexed part 3-10 mm. broad. 




[Vol. 7 
132 ANNALS OF THE MISSOUBI BOTANICAL GARDEN 



On rotting logs and limbs of frondose species such as Acer, 
Betula, Fagus, Quercus, and Tilia. Canada to West Indies and 
westward to British Columbia. April to October in the north 
and October to March in the West Indies. Common. Occurs 
in Scandinavia also. 

The specimens upon which were based the original descrip- 
tions of S. Murrayi and its synonyms were resupinate; in each 
instance the species was included in Stereum or Thelephora, 
although longitudinally arranged hyphae are not present and 
do not constitute an intermediate layer. The distinguishing 
characters of the resupinate specimens are their thickness, 
pallid to pale avellaneous color, tubercular and deeply cracked 
hymenium, abundance of vesicular organs throughout the whole 
thickness of the fructification, and occurrence on a frondose 
substratum. The horny crust forming the upper side of the 
pileus is similar to that of some species of Fomes and is unique 
among our Stereums, but the reflexed stage is so rare that this 
character does not often afford help in recognizing the species. 
The geographical distribution in three widely separated areas is 
remarkable; it seems probable that the European stations in 
Norway and Sweden should be regarded as merely outlying 
stations of a common North American species; it is very strange 
that a species presumably northern should be well established 
in Cuba and Jamaica and absent from Florida and the Carolinas, 
yet specimens from all three isolated regions are identical in 
aspect and microscopical structure. 

Specimens examined: 
Exsiccati: Ell. & Ev., Fungi Col., 704, under the name Stereum 

rugosum; Ell. & Ev., N. Am. Fungi, 2903, under the name 
Corticium colliculo sum ; Shear, N. Y. Fungi, 51, under the 
name Stereum rugosum. 

Norway: M. N . Blytt, type of Stereum tuberculosum (in Herb. 

Fries) . 
Sweden: Island of Gotland, on Abies excelsa, L. Romell, 135. 
Canada: J. Macoun, 18, 43, 60; Billings Bridge, J. Macoun, 44; 

Lower St. Lawrence Valley, J. Macoun, 69, 72. 
Ontario: J. Deamess, 1022 (in Mo. Bot, Gard. Herb., 22682); 

Blackwater, J. McFarlane, Univ. Toronto Herb., 330 (in 

Mo. Bot. Gard. Herb., 44865); Harraby, Lake Rosseau, 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 133 



E. T. & S. A. Harper, 730; London,,/. Dearness, two 
collections, and in Ell. & Ev., Fungi Col., 704; Ottawa, 
J. Macoun, 12, and 676 — the latter comm. by W. G. Farlow 
(in Mo. Bot. Gard. Herb., 56757) ; Toronto, Algonquin Park 
and Lome Park, J. H. Fault, Univ. Toronto Herb., 500 
and 333 respectively (in Mo. Bot. Gard. Herb., 44854 and 
44873). 

Maine: F. L. Harvey, comm. by P. L. Ricker, and F. L. Harvey, 

type of Stereum pulverulentum (in N. Y. State Mus. Herb.) 

and cotype comm. by P. L. Ricker; Portage, L. W. Riddle, 

19; Sebec Lake, W. A. Murrill, 2304 (in N. Y. Bot. Gard. 

Herb, and Mo. Bot. Gard. Herb., 56755). 
New Hampshire: Chocorua, W. G. Farlow; Crawford Notch, 

L. 0. Overholts, 4582 (in Mo. Bot. Gard. Herb., 55640) ; 

Groton, J. Blake, comm. by P. L. Ricker; Hebron, P. 

Wilson (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. 

Herb., 56756) ; Shelburne, W . G. Farlow (in Farlow Herb.). 
Vermont: Bristol, E. A. Burt; Grand View Mt., E. A. Burt, 

two collections; Middlebury, E. A. Burt, two collections; 

Ripton, E. A. Burt, two collections and also near Abby 

Pond and Lost Pleiad Lake. 
Massachusetts: Murray, comm. by Sprague, 546, authentic 

specimen of Thelephora Murrayi (in Curtis Herb., 5809). 
New York: Alcove, C. L. Shear, 1206, 1311, and in Shear, N. Y. 

Fungi, 51; Altamont, E. A. Burt; Floodwood, E. A. Burt; 

Fulton Center, L. M. Underwood (in N. Y. Bot. Gard. Herb. 

and Mo. Bot. Gard. Herb., 56274); Horicon, C. H. Peck 

(in N. Y. State Mus. Herb, and Mo. Bot. Gard. Herb., 

56107); Ithaca, C. J. Humphrey, 549; Lake Placid, W. A. 

& E. L. Murrill, 194 (in N. Y. Bot. Gard. Herb, and Mo. 

Bot. Gard. Herb., 56756); North Elba, C. H. Peck, 1; 

Seventh Lake, Adirondack Mts., B. M. Duggar & F. C. 

Stewart; West Ann, S. H. Burnham, 4 (in Mo. Bot. Gard. 

Herb., 43997). 

West Virginia: Nuttallburg, L. W. Nuttall, in Ell. & Ev., Fungi 

Col., 704. 

Michigan: Houghton, C. H. Kaufman, comm. by N. Y. State 

Mus. Herb, (in Mo. Bot. Gard. Herb., 55812); Sailors' 
Encampment, Miss A. 0. Stucki, 5; Vermilion, A. H.W. 
Povah, 190 (in Mo. Bot. Gard. Herb., 17615). 



[Vol,. 7 



134 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Wisconsin: Ladysmith, C. J. Humphrey, 1914 (in Mo. Bot. 

Gard. Herb., 42916). 
Idaho: Priest River, J. R. Weir, 362, 379 (in Mo. Bot. Gard. 

Herb., 16533, 17115). 
British Columbia: Agassiz, J. R. Weir, 351 (in Mo. Bot. Gard. 

Herb., 8066). 
Cuba: C. Wright, 269, type (in Kew Herb, and Curtis Herb.); 

Alto Cedro, Earle & Murrill, 491, comm. by N. Y. Bot. 

Gard. Herb.; Ciego de Avila, Earle & Murrill, 590, comm. 

by N. Y. Bot. Gard. Herb.; Herradura, Earle & Murrill, 

188, comm. by N. Y. Bot. Gard. Herb. 
Porto Rico: Rio Piedras, /. A. Stevenson, 

Gard. Herb., 7584). 
Jamaica: Constant Spring Hotel grounds, W. A. & E. L. 

Murrill, 34, comm. by N. Y. Bot. Gard. Herb. ; New Haven 

Gap, W. A. & E. L. Murrill, 771, comm. by N. Y. Bot. 

Gard. Herb.; Port Antonio, F. S. Earle, 575, comm. by 




(in Mo. Bot. 



N. Y. Bot. Gard. Herb. 



36. S. saxitas Burt, n. sp. 



Plate 4, fig. 33. 



Type: in Mo. Bot. Gard. Herb, and N. Y. Bot. Gard. Herb. 
Fructification thick, stratose, stony-hard throughout, resupi- 
nate, effused, becoming narrowly reflexed, the reflexed portion 

black above, irregular, stony; 
hymenium even or tubercular, 
not shining, drying cartridge- 
buff to whitish; in structure 1-5 
mm. thick, stratose, composed 



mm 



111! 





s 



of alternating pale and darker 
layers but with a horn-like 
translucent luster throughout 
when cut; a few vesicular or- 
gans 20-25x12-15 n present 
along the under portion of each 
stratum; no cystidia; spores 
hyaline, even, 4-5 X 3-4 n. 
Resupinate portion 3-6 cm. in 

diameter; reflexed margin 2-4 mm. broad. 

On bark of apparently a frondose species. Mexico and 

Jamaica. December and May. 



mi 

Fig. 16. S. saxitas. Section of hy- 
menial region X 90, showing vesicular 
bodies; spores, s, X 665. 






1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



135 



S. saxitas resembles in aspect S. Murrayi, and relationship tc 
this species is further shown by the presence of vesicular organs 
however, it is thicker than S. Murrayi, stony-hard throughout 



but few vesicular cells, and has subglobose spore 
! is so extremely hard that it has been possible 



Its 



sections for microscopic details of only the hymenium and 
nearly adjacent regions even after prolonged soaking in water. 

Specimens examined: 
Mexico : Guernavaca, W. A.& E.L. Murrill, If.19, type, comm. by 

N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 54552). 
Jamaica: John Crow Peak, D. S. Johnson (in N. Y. Bot. Gard. 

Herb.. Mo. Bot. Gard. Herb., 56758, and Burt Herb.). 



37. S. styracifluum Schweinitz, Naturforsch. Ges. Leipzig 
Schrift. 1: 105. 1822 (under B. Sterea of Thelephora); Fries, 
Epicr. 549. 1838; Sacc. Syll. Fung. 6: 569. 1888. 

Plate 4, figs. 34, 35. 

Thelephora styraciflua Schweinitz in Fries, Elenchus Fung. 

1: 177. 1828; Schweinitz, Am. Phil. Soc. Trans. N. S. 4: 167. 

1832. 

Type: in Herb. Schweinitz and portions in Herb. Fries and 

Curtis Herb. 

Fructification coriaceous, resupinate and effused, with a nar- 
row, free marginal portion, or slightly reflexed, tomentose, dry- 
ing pinkish buff to cinnamon-buff; 
hymenium dull, pruinose, not mul- 
tizonate, drying pinkish buff, exud- 
ing a yellow milk when compressed 
and becoming dark-discolored, con- 
tracting in drying and splitting; in 



700 



thick 



intermediate layer bordered 1 
upper side by a pale golden 
not denser than the rest 




rest of the 
very densely 
3 n in diam- 
with pale-colored conducting 



layer, composed of 
arranged hyphae 2\ 



Fig. 17. S. styracifluum. Section 
of hymenial region X 488, showing 
conducting organs. From type. 



diameter which curve into the hymenium 



ystidia; spores hyaline, even, slightly curved 



3 



v 



1 9 A IV ° L - 7 

lOO ANNALS OP THE MISSOURI BOTANICAL GARDEN 



Resupinate portion 3X2 cm.; the free margin up to 5 mm. 
broad. 

On under side of dead, fallen limbs of Liquidambar and mossy 
dead trunk of Carpinus. North Carolina and Alabama. Jan- 
uary. Rare. 

S. styracifluum is intermediate between S. rameale and S. 
rugosum; in the region where it occurs it is likely to be regarded 
as a resupinate form of S. rameale, from which it differs in darker 
and more irregular hymenial surface, greater thickness of fructi- 
fication, margin sometimes with a black edge, and reflexed part 
tomentose to the margin; the pale-colored conducting organs 
are similar in the two species but rather more abundant in S. 
styracifluum. The general aspect is so similar to that of S. 
rugosum, very common in Europe, that the yellow milk of S. 
styracifluum was properly regarded by Schweinitz as an impor- 
tant distinctive character of the American species; other differ- 
ences are that the intermediate layer is much broader and denser 
than that of S. rugosum, that the hymenium is only 20-30 ft 
broad, never zonate, and that the conducting organs are much 
less numerous and paler than in S. rugosum. 

Specimens examined: 
North Carolina: Salem, Schweinitz, type (in Schweinitz Herb., 

Fries Herb., and Curtis Herb.). 
Alabama: Auburn, on Carpinus, F. S. Earle & C. F. Baker (in 

Burt Herb, and Mo. Bot. Gard. Herb., 5061). 

38. S. gausapatum Fries, Hym. Eur. 638. 1874; Sacc. Syll. 
Fung. 6: 560. 1888; Massee, Linn. Soc. Bot. Jour. 27: 180. 
1890. Plate 4, fig. 36. 

Thelephora gausapata Fries, Elenchus Fung. 1: 171. 1828; 
Epicr. 538. 1838.— T. spadicea Fries, Elenchus Fung. 1: 176. 
1828 (not T. spadicea Persoon, Syn. Fung. 568. 1801. See 
Bresadola, I. R. Accad. Agiati Atti III. 3: 106. 1897). 
Stereum spadiceum Fries, Epicr. 549. 1838; Hym. Eur. 640. 
1874; Berkeley, Outlines Brit. Fung. 270. 1860; also of more 
recent English authors. — S. spadiceum var. plicatum Peck, N. Y. 
State Mus. Rept. 50 : 132. 1897.— £. cristulatum Quelet, Champ. 
Jura et Vosges 3: 15. pi. l.f. 15. 1875.— S. occidentale Lloyd, 
My c. Writ. 5. Letter 69:12. 1919. 

Type: specimen from Fries in Kew Herb. 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 137 




Fructifications coriaceous, effuso-reflexecl or somewhat dimid 
iate, usually cespitose-imbricated, confluent, varying from vil 
lose to hirsute, buckthorn-brown, more 
or less radially plicate; hymenium bleed- 
ing when fresh if cut or bruised, drying 
snuff-brown and more or less darker dis- 
colored; in structure 600-700 n thick 
exclusive of the hairy covering, com- Fi e- 18 - s - gausapatum. 
posed of densely and longitudinally ar- ?^ on ° f hy^enial region 

, . , > , n i , X 68, showing distribution 

ranged hyphae, with flexuous, colored of conducting organs, 
conducting organs 75-120x5 ft, very 

numerous in the hymenium; no cystidia; spores hyaline, even, 
5— 8X2|— 3§ ju. 

Singly or covering areas up to 10 cm. and more in diameter; 
reflexed portion about 1 cm. broad, 1-2 } cm. long or more, or 
with small pilei or lobes 1-1 1 cm. in diameter. 

On stumps of Quercus usually. Canada to Alabama and 
westward to Washington and California. August to March. 
Common. 

S. gausapatum is usually recognizable at sight by its clustered 
fructifications tobacco-colored above and clothed with a heavy 
villose or strigose coat, by the rather dark hymenium which 
bleeds when cut and becomes somewhat darker discolored in 
drying, and by the occurrence on oak. Sectional preparations 
show very numerous, colored conducting organs in the hyme- 
nium. S. australe of the Gulf states bleeds and has colored 
conducting organs, although fewer, but its fructifications do not 



form dense clusters and 



S 



lentum has the same geographical distribution as S. gausapatum 
and bleeds when fresh and has colored conducting organs, 
but has small fructifications occurring on conifers only. The 
hairy covering of the pileus is greedily devoured by herbarium 
insects, leaving the pilei bare of their normal covering if speci- 
mens are not protected against their depredations, but, except 
for insect depredation, this covering is a persistent character. 

Fries described the effuso-reflexed stage of S. gausapatum 
under the name T. spadicea, confusing this stage with the more 
southern and specifically different Thelepkora spadicea of Per- 
soon, which does not occur in America. It seems preferable 



[Vol. 7 



138 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



to use the name S. gausapatum for our species, although unfor- 
tunately the other name is in general use in England, and leave 
the name S. spadiceum available for use in its original sense as 
continental mycologists do. It is surprising that specimens of 
S. gausapatum do not occur in Herb. Schweinitz under some 

name or other. 

Specimens examined: 

Exsiccati: Bartholomew, Fungi Col., 2883, 4292; Berkeley, 

Brit. Fungi, 144; Cooke, Fungi Brit., 107; Ellis, N. Am. 
Fungi, 325; Ell. & Ev., N. Am. Fungi, 3413, under name 
Stereum hirsutum; Ell. & Ev., Fungi Col., 218; Ravenel, 
Fungi Car. 2: 32; Fungi Am., 447; Romell, Fungi Scand. 
Exs., 28, 122. 

Sweden: Stockholm, L. Romell, J+5, 1+6, 238, and in Romell, 

Fungi Scand. Exs., 28, 122. 

England: M. J. Berkeley, in Berkeley, Brit. Fungi, 144; Epping, 

M. C. Cooke, in Cooke, Fungi Brit., 107. 
Holland: Amsterdam, C. A. J. A. Oudemans, in Oudemans, 

Fungi Neerland., 239 (in Mo. Bot. Gard. Herb.). 
France: authentic specimen of Stereum cristulatum from Quelet 

(in Herb. Fries); wall of German trench, Lieut. G. W. 

Martin, comm. by P. J. Anderson, 3 (in Mo. Bot. Gard. 

Herb., 55848) ; St. Sernin, Aveyron, A. Galzin, 1265, comm. 

by H. Bourdot, 16234; Corrombles, F. Fautrey, from Lloyd 

Herb., 3312. 
Italy: Trentino, G. Bresadola. 
Canada: Carleton Place, J. Macoun, J^19. 
Ontario: Lake Joseph, T. Langton, Univ. Toronto Herb., 590 

(in Mo. Bot. Gard. Herb., 44846); London, J. Dearness; 

Swansea, /. H. Fault, Univ. Toronto Herb., 375 (in Mo. 

Bot. Gard. Herb., 44931); Toronto, J. H. Fault, G. H. 

Graham, T. Langton, R. P. Wodehouse, Univ. Toronto 

Herb., 372, 376, 676, 679, 591, 597, 368 (in Mo. Bot. Gard. 

Herb., 44946, 44932, 44923, 44935, 44849, 44840, 44855, 

respectively). 
Vermont: Lake Dunmore, E. A. Burt, three collections; Mid- 

dlebury, E. A. Burt. 
Massachusetts: Mt. Auburn, E. A. Burt; Stoneham, C. L. 

Shear, 1283; Way land, A. B. Seymour, TS6 (in Mo. Bot. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 139 



Gard. Herb., 13939); Waverly, G. R. Lyman, 121; Weston, 
A. B. Seymour, T10 (in Mo. Bot. Gard. Herb., 19621). 
Connecticut : West Hartford, C. C. Hanmer, 2670 (in Mo. Bot. 

Gard. Herb., 42605). 
New York: Sartwell (in Mo. Bot. Gard. Herb., 5046, 5102); 

Cold Spring Harbor, H. J. Banker (in Mo. Bot. Gard. 
Herb., 54434); Green Lake, P. Wilson, 52 (in Mo. Bot. 
Gard. Herb., 54745); Ithaca, G. F. Atkinson, 223 0. S., 
2140, 7986, 7986b, H. H., 5088, C. J. Humphrey, F. A. 
Wolf, 22948; N. Greenbush, C. H. Peck, in Ellis, N. Am. 
Fungi, 325; Poughkeepsie, W. R. Gerard, 271 (in N. Y. 
Bot. Gard. Herb.); Shakers, S. H. Burnham, 16 (in Mo. 
Bot. Gard. Herb., 44010); St. Regis Falls, L. A. Zimm, 94 
(in Mo. Bot. Gard. Herb., 21941); Williamsbridge, P. 
Wilson, 2 (in Mo. Bot. Gard. Herb., 54746); White Plains, 
L. M. Underwood (in N. Y. Bot. Gard. Herb, and Mo. Bot. 

Gard. Herb., 56700). 
New Jersey: Newfield, J. B. Ellis, in Ell. & Ev., Fungi Col., 218. 
Pennsylvania: Kittanning, D. R. Sumstine, 5, 6, 8; Spruce 

Creek, /. H. Faull, Univ. Toronto Herb., 371, 672 (in Mo. 

Bot. Gard. Herb., 44925, 44938); Trexlertown, C. G. Lloyd, 

0054. 
Delaware : Newark, H. S. Jackson. 

Maryland: Takoma Park, C. L. Shear, 1018, 1201, 1270, 1273. 
Virginia: Clarendon, W. H. Long, 12617 (in Mo. Bot. Gard. 

Herb., 55103); Park Lane, W. H. Long, 12860 (in Mo. 

Bot. Gard. Herb., 55109). 
North Carolina: Biltmore, C. Harrison, comm. by P. L. Ricker, 

E. Bartholomew, 5663 (in Mo. Bot. Gard. Herb., 44262); 

Blowing Rock, G. F. Atkinson, 4318, 4328; Chapel Hill, 

W. C. Coker, 834, $821 (in Mo. Bot. Gard. Herb., 56670, 

56671). 
South Carolina: Aiken, H. W. Ravenel, in Ravenel, Fungi Am., 

447; Black Oak, H. W. Ravenel, in Ravenel, Fungi Car. 

2: 32. 

Georgia: Tallulah Falls, A. B. Seymour, comm. by W. G. Far- 
low, C. C. (in Mo. Bot. Gard. Herb., 44604). 

Alabama: Auburn, F. S. Earle (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56287, 56703), and C. F. Baker, 



*At\ [VoL - 7 

14U ANNALS OF THE MISSOURI BOTANICAL GARDEN 



50 (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 

56702); Montgomery Co., R. P. Burke, 24, 38 (in Mo. Bot. 

Gard. Herb., 17651, 4925). 
Louisiana: St. Martinville, A. B. Langlois, 165. 
Michigan: Beat, 57, comm. by N. Y. State Mus. Herb, (in Mo. 

Bot. Gard. Herb., 55810); Ann Arbor, C. H. Kauffman, 

37 (in Mo. Bot. Gard. Herb., 18995); Glen Lake, C. G. 

Lloyd, 02551 . 
Ohio: Cincinnati, C. G. Lloyd, 02820; College Hill, W. Holden, 

comm. by Lloyd Herb. 
Indiana: Millers, E. T. & S. A. Harper, 678. 
Illinois: River Forest, E. T. & S. A. Harper, 708; Riverside, 

E. T.& S. A. Harper, 686. 

West Virginia: Nuttallburg, L. W. Nuttall, in Ell. & Ev., N. 

Am. Fungi, 3413. 

Kentucky: S. A. Price (in Mo. Bot. Gard. Herb., 5136). 
Wisconsin: Madison, E. T. & S. A. Harper, 942, Miss A. D. 

Stucki, 32, and W. Trelease, 84 (in Mo. Bot. Gard. Herb., 
5101). 

Iowa: Webster Co., 0. M. Oleson, 2, 3, 5. 

Missouri: Columbia, B. M. Duggar, 358, 392, 573; St. Louis, 

C. R. Ball & H. H. Hume, and E. A. Burt (in Mo. Bot. 

Gard. Herb., 5023, 21989). 
Arkansas: Fayetteville, E. Bartholomew, in Bartholomew, Fungi 

Col., 2883; Womble, W. H. Long, 19849 in part (in Mo. 

Bot. Gard. Herb., 20271). 

Texas: Joaquin, E. Bartholomew, in Bartholomew, Fungi Col., 

4292. 

Nebraska: Lincoln, C. L. Shear, 1017 ; Roco, C. L. Shear, 1012. 

Kansas: Bourbon Co., A. G. Garrett, 86, 129. 

British Columbia: Kootenai Mts., near Salmo, J. R. Weir, 502 

(in Mo. Bot. Gard. Herb., 21630). 
Washington: Seattle, S. M.Zeller, 109 (in Mo. Bot. Gard. Herb., 

44142); T. C. Frye, 2007 (in N. Y. Bot. Gard. Herb.); 

Whidley Is., N. L. Gardner, Univ. Calif. Herb., 1033 (in 

Mo. Bot, Gard. Herb., 44151). 
Oregon: Corvallis, C. E. Owens, 2085 (in Mo. Bot. Gard. Herb., 

44247), W. A. Murrill, 903, comm. by N. Y. Bot. Gard. 

Herb, (in Mo. Bot. Gard. Herb., 55720); Portland, /. R. 

Weir, 396 (in Mo. Bot. Gard. Herb., 14094). 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 141 



California: I. M. Johnston, comm. by C. G. Lloyd, part of type 

of Stereum occidentale (in Mo. Bot. Gard. Herb., 56762); 

Alameda Co., L. S. Smith, Univ. Calif. Herb., 403 (in Mo. 

Bot. Gard. Herb., 44150); Preston's Ravine, Palo Alto, W. 

A. Murrill & L. S. Abrams, 1190, comm. by N. Y. Bot. 

Gard. Herb, (in Mo. Bot. Gard. Herb., 55711); Redwood 

Park, W. H. Long, 12604 (in Mo. Bot. Gard. Herb., 55100); 

Santa Barbara, O. M. Oleson, 7, 15. 
Arizona: C. G. Pringle, comm. by W. G. Farlow. 
Mexico: San Luis Potosi, comm. by U. S. Dept. Agr. Herb. 



39. S. australe Lloyd, Myc. Writ. 4. Letter 48: 10. 1913; 
and ibid. Letter 60: 15. 1916. Plate 4, fig. 37. 

An Thelephora mytilina Fries, Elenchus Fung. 1 : 175. 1828? 

Type: in Lloyd Herb, and Mo. Bot. Gard. Herb. 

Fructification coriaceous, attached by the resupinate side and 
umbo, broadly reflexed, sometimes laterally confluent, densely 
tomentose, becoming concentrically furrowed and very rarely 
glabrous and showing the shining chestnut surface of the pileus 
in one or more of the furrows, the margin entire, sometimes 
becoming blackish; hymenium even, glabrous, drab-gray to avel- 
laneous, becoming red-discolored where cut or bruised, and 
sometimes bleeding; in structure 900 n thick, composed of dense- 
ly, longitudinally arranged hyphae, among which are a few col- 
ored conducting organs 3§-4£ n in diameter which curve into the 
hymenium between the basidia; no cystidia nor gloeocystidia 
present; spores hyaline, even, flattened on one side, 4^|X2^-3 n, 
few found. 

Fructifications with resupinate portion 1--3 cm. broad, reflexed 
portion 1-4 cm. broad, 1-5 cm. long and sometimes more by 
lateral confluence. 

On hardwood logs. Florida and Mississippi to Brazil. Au- 
gust to December in the north and in July in Brazil. Appar- 
ently rare. 

Stereum australe combines the characters of S. fasciatum and 
S. gausapatum. Its general aspect resembles that of specimens 
of S. fasciatum in a middle period of development when they are 
effuso-reflexed and have the umbo developed, but the specimens 
of S. australe have a broader resupinate portion than those of S. 



[Vol. 7 
142 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



fasciatum and are not wedge-shaped and attached merely by 
the umbo in any specimens which I have seen; the bleeding or 
red-discoloration of the hymenium when cut or bruised and the 
presence of colored conducting organs are additional characters 
which separate S. australe from S. fasciatum. S. australe may 
be distinguished from S. gausapatum by not having its reflexed 
portion crisped nor consisting of small pilei which stand out near 
together and in imbricate arrangement from a common resupi- 
nate portion. 

In case of the collection from Mississippi, it was noted that the 
substratum was badly sap-rotted. 

If original specimens of Thelephora mytilina, collected by Lund 
in Brazil, are still in existence, I believe that they will be found 
cospecific with S. australe. The geographical range of S. australe 
and the description of T. mytilina favor this belief. Fries's de- 
scription was probably based on dried specimens, and it does not 
mention bleeding of the hymenium nor such a microscopical 
character as colored conducting organs, for such a microscopic 
detail was not noted in those days, but the blackening of the 
edge of the pileus which was observed by Fries is an indication 
of a bleeding hymenium and colored conducting organs. 

Specimens examined: 
Florida: type comm. by C. G. Lloyd (in Mo. Bot. Gard. Herb., 

56608); Kissimme, C. J. Humphrey, 3532 (in Mo. Bot. 

Gard. Herb., 3370). 
Mississippi: Laurel, C. J. Humphrey, 5434- 
Mexico: Jalapa, W. A. & E. L. Murrill, 189, comm. by N. Y. 

Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 54446). 
Canal Zone: Gatun, M. A. H. (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56707). 
Grenada: W. E. Broadway (in N. Y. Bot. Gard. Herb, and Mo. 

Bot. Gard. Herb., 56625, 56626). 
Venezuela: Caracas, Mr. & Mrs. J. N. Rose, 22038 (in N. Y. 

Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 56657). 
Brazil: Rio de Janeiro, J. N. Rose & P. G. Russell, 21480 (in 

N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 56627). 



40. S. rugosum Persoon, Roemer Neues Mag. Bot. 1: 110. 
1794; Fries, Epicr. 552. 1838; Myc. Eur. 643. 1874; Berk- 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



143 



eley, Brit. Fungi, 271. 1860; Sacc. Syll. Fung. 6: 572. 



1888; 



Massee, Linn. Soc. Bot. Jour. 27: 191. 



1890. 



Plate 4, figs. 38, 39. 

Thelephora rugosa Persoon, Syn. Fung. 569. 1801; Myc. 
Eur. 1: 127. 1822; Albertini & Schweinitz, Consp. Fung. 274. 
1805; Fries, Syst. Myc. 1: 439. 1821; Elenchus Fung. 1: 177. 
1828. 

Illustrations: Istvanffi, Jahrbuch. f. wiss. Bot. 29: pi. 4* 
f. 11; pi 5.f. 19. 

Fructifications coriaceous-corky, usually resupinate and 
effused, with a narrow, free, marginal portion, or sometimes 
reflexed, silky at first and pinkish buff, at 
length concentrically furrowed, radially 
pitted and weathering gray, the margin 
thick, entire; hymenium dull, pruinose, 
drying pinkish buff to drab-gray, when 



fresh bleeding where wounded; in struc- 
ture 500-1800 n thick, with the interme- 
diate layer bordered on the upper side by 
a dense golden zone and on the lower side 
by a two- to many-zoned hymenial layer 



mm 




1/ 




Fig. 19. S. rugosum. 
Section X 19; interme- 
diate layer, i; dense 



120-1200 m thick, hyphae of intermediate golden zone, z; the scat- 



layer 



L 2 



3 n in diameter; dark-colored 



tered darker lines in 
hymenial zones show 
distribution of conduct- 
ing organs. 



conducting organs very numerous, 3-6 /x 
in diameter; no cystidia; spores hyaline, 
even, flattened on one side, 7-10X3-4 //. 

Resupinate on areas 2-6 cm. in diameter; free or reflexed 
margin 2-12 mm. broad. 

On stumps of Alnus, Corylus, Quercus, Betula, and other fron- 
dose species. Newfoundland, Ontario, New York, and moun- 
tains of North Carolina. July to October. Rare in North 
America, common in Europe. 

Although usually resupinate and likely to be regarded as a 
Corticium by collectors, nevertheless sectional preparations 
show the highly developed characteristic structure of a Stereum, 
with intermediate layer of longitudinally arranged hyphae, 
golden crust, etc. The bleeding of the hymenium and the abun- 
dant colored conducting organs locate the species among the 
Stereums in the group with S. gausapatum, S. australe, and S. 



[Vol. 7 

144 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



sanguinolentum , from each of which S. rugosum is sharply distinct 
by its two- to several-zoned hymenium — a character by which 
the species is also separable in dried herbarium condition from 
S. styracifluum when no observations have been recorded as to 
the color of the milk of specimens in fresh condition. 

Specimens examined: 
Exsiccati: Berkeley, Brit. Fungi, 145; Krieger, Fungi Sax., 1853, 

1853b; Rabenhorst, Herb. Myc, 503; Romell, Fungi 
Scand. Exs., 30; de Thumen, Myc. Univ., 1009. — All speci- 
mens distributed as S. rugosum in American exsiccati were 
misdetermined. 
England: M. J. Berkeley, in Berkeley, Brit. Fungi, 145; Epping 

Forest, E. A. Burt; Kew Garden, G. Massee. 
Sweden: L. Romell, lfi-J+2; Femsjo, E. A. Burt; Stockholm, 

L. Romell, in Romell, Fungi Scand. Exs., 30; Upsala, 
E. P. Fries (in Curtis Herb.). 
Finland: Mustiala, P. A. Karsten, in de Thumen, Myc. Univ., 

1007. 

Germany: Dresden, in Rabenhorst, Herb. Myc, 503; Saxony, 

Uttewalder Grunde, W. Krieger, in Krieger, Fungi Sax., 
1853, 1853b. 

Hungary: Tatra Magna, Locse, V. Greschik, comm. by G. 

Bresadola. 
Italy: Trentino, G. Bresadola, two collections. 
France: Allier, St. Priest, H. Bourdot, 15028. 
Newfoundland: Bay of Islands, A. C. Waghorne, 160 (in Mo. 

Bot, Gard. Herb., 5096); Trinity Bay, A. C. Waghorne, 

1 (in Mo. Bot. Gard. Herb., 5098). 
Quebec: Gaspe, J. Macoun, and 254 (in N. Y. State Mus. Herb. 

and Mo. Bot, Gard. Herb., 56094). 
Ontario: Ottawa, J. Macoun, 38. 
New York: Fall Creek, G. F. Atkinson, 949. 
North Carolina: Blowing Rock, G. F. Atkinson, 4189. 

41. S. sanguinolentum Albertini & Schweinitz, Consp. Fung 
274. 1805 (under B. Sterea of Thelephora) ; Schweinitz, Natur- 
forsch. Ges. Leipzig Schrift. i: 106. 1822; Fries, Epicr. 549. 
1838; Hym. Eur. 640. 1874; Berkeley, Brit. Fungi, 271. 1860; 
Sacc. Syll. Fung. 6: 564. 1888; Massee, Linn. Soc. Bot. Jour. 



7: 189. 1890. 



Plate 5, fig. 40. 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



145 



Thelephora sanguinolenta Alb. & Schw. in Fries, Syst. Myc 
i:440. 1821; Elenchus Fung, i: 178. 1828.— Stereum bal 

99. 1875; ibid 

1888; Massee. Linn 



eum 



Peck, N. Y. State Mus. Rept. 27 



75. 1879; Sacc. Syll. Fung. 6 



Soc. Bot. Jour. 27 



1890. — S. balsameum form reflexum 



Peck, N. Y. State Mus. Rept. 47: 1. 
ten, Finska Vet.-Soc. Bidrag Natur 



1894. — S. rioens Kars 



bid. 48 



Folk 37 



1889; Sacc. Syll. Fung 



1882; 



1895. 



Illustrations: Gillet, Hymenomycetes ; Greville, Crypt 



pi. 225; Istvanffi, Jahrbiich. f. wiss. Bot. 29: 



Klotzsch 



Dietrich, Fl. R 



2nded. pl.28.f 



Patouillard, Tab. Anal./. 



4- /• 7-10 

Nees, Syst 



Fructifications coriaceous, thin, effused, and reflexed, with 
>per surface villose to silky and the hairs appressed and some- 
lat radiately arranged, dry- 
z pinkish buff to pale olive- 



buff, the margin thin 



hy 



menium glabrous, bleeding 
where wounded, contracting 
in drying and cracking to the 
substratum in the resupinate 
portion, drying avellaneous 
to wood-brown: in structure 







^ 




3 



Fig. 20. S. sanguirwlentum. Section 
hymenial 



400-600 fx thick, with inter- tionof conducting organs; spores, s, X 488. 

mediate layer bordered on 

side by a narrow, dense golden zone, and composed 

aline hyphae 3 m in diameter and of 



densely arranged h^ 



hich curve into the 



hymenium and are usually numerous there; no cystidia 
white in spore collection, even, slightly curved, 6-7 X2h 



1-5 



cm 



diamet 



2 M- 

reflexed mar 



Resupinate portions 
2-10 mm. broad. 

On stumps and logs of Pinus, Abies, and Tsvga. Ontario to 
Pennsylvania and westward to British Columbia and California. 
July to March. Frequent. 

S. sanguinolentvm is commonly resupinate or barely reflexed, 
so that it is best recognized by its occurrence on conifers and 
bleeding of the hymenium where wounded, or becoming merelv 



d-discolored along the ed 



d if 



wound 



£ 



[Vol. 7 



146 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



made during dry weather. The somewhat drab color the hy- 
menium assumes in drying and its deep cracks are highly char- 
acteristic of dried specimens. Colored conducting organs are 
abundant in the hymenium and subhymenium and should 
be demonstrated if other characters leave the determination 

doubtful. 

Specimens examined: 

Exsiccati: Krieger, Fungi Sax., 160; Romell, Fungi Scand. 

Exs., 29; de Thumen, Myc. Univ., 2010, and 2111, under 
the name Stereum rigens. 

Sweden: L. Romell, 43, 44; Lapland, L. Romell, 401 bis; Stock- 
holm, L. Romell, in Romell, Fungi Scand. Exs., 29; Upsala, 

E. A. Burt. 
Finland: Mustiala, P. A. Karsten, in de Thumen, Myc. Univ. 

2010,2111. 
France: Allier, H. Bourdot, 5586, 7591. 
Italy: G. Bresadola; Florence, G. Arcangeli (in Mo. Bot. Gard. 

Herb., 44565). 
Newfoundland: Bay of Islands, A. C. Waghorne, 837, 350, the 

latter determined by Peck as S. balsameum (in Mo. Bot. 

Gard. Herb., 5099, 5056). 
Canada: comm. by J. B. Ellis, 5070 (in Kew Herb., under the 

name Stereum triste as determined by Cooke). 
Quebec: Montreal, R. J. Blair, comm. by L. O. Overholts, 

3787, 4107 (in Mo. Bot. Gard. Herb., 55097, 55638). 
Ontario: Bond Lake, J. H. Fault, Univ. Toronto Herb., 320 

(in Mo. Bot. Gard. Herb., 44875); Casselman, J. Macoun, 

859; Lake Nipegon, J. Macoun, 103; Ottawa, J. Macoun, 

11; Toronto, R. P. Wodehouse, Univ. Toronto Herb., 369 

(in Mo. Bot. Gard. Herb., 44850); York Mills, J. H. Fault, 

Univ. Toronto Herb., 318 (in Mo. Bot. Gard. Herb., 44877). 
Maine: Piscataquis Co., W. A. Murrill, 2029 (in N. Y. Bot. 

Gard. Herb, and Mo. Bot. Gard. Herb., 56705); Portage, 

L. W. Riddle, 18. 
New Hampshire: Chocorua, W. G. Farlow, 4', Tuckerman's 

Ravine, Mt. Washington, L. O. Overholts, 4949 (in Mo. 

Bot. Gard. Herb., 56343). 
Vermont: Little Notch, Middlebury, and Ripton, E. A. Burt. 
Massachusetts: R. J. Blair, 827, comm. by L. O. Overholts, 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 147 

4118 (in Mo. Bot. Gard. Herb., 55641), and D. W. Weis, 
comm. by C. G. Lloyd, 129 (in Mo. Bot. Gard. Herb., 

56708). 

New York: Adirondack Mts., C. H. Peck, type of Stereum bal- 

sameum (in N. Y. State Mus. Herb.) ; Alcove, C. L. Shear, 
1136; Cayuga Lake Basin, G. F. Atkinson J, 3028, 8271, and 
H. Hasselbring, 3408; Glasco, P. Wilson, 38 (in Mo. Bot. 
Gard. Herb., 54743); Ithaca, C. J. Humphrey, 805. 

Pennsylvania: Shingleton Gap, A. S. Rhoads, 9 (in Mo. Bot. 

Gard. Herb., 44086). 

North Carolina: Salem, Schweinitz (in Herb. Schweinitz). 

Michigan: Gogebic Co., E. A. Bessey, 224 (in Mo. Bot. Gard. 

Herb., 56563). 

* 

Montana: Anaconda, /. R. Weir, 11973 (in Mo. Bot. Gard. 



Herb., 56727); Elkhorn, J. R. Weir, 9749 (in Mo. Bot. 
Gard. Herb., 56224); Evaro, J. R. Weir, 413 (in Mo. Bot. 
Gard. Herb., 14773). 

Colorado: Ouray, C. L. Shear, 1186. 

New Mexico: Sandia Mts., W. H. Long, 21576, 21597 (in Mo. 

Bot. Gard. Herb., 55154, 55116); Tyom Experiment Sta- 
tion, W. H. Long, 2155^ (in Mo. Bot. Gard. Herb., 55115). 

Idaho: Priest River, J. R. Weir, 47, 347 (the latter in Mo. Bot. 

Gard. Herb., 9989); Sandpoint, E. E. Hubert, comm. by 
J. R. Weir, 11612 (in Mo. Bot. Gard. Herb., 56726). 

British Columbia: Agassiz, J. R. Weir, 387 (in Mo. Bot. Gard. 

Herb., 20887); Hastings, /. Macoun, 27; Kootenai Mts., 

near Salmo, J. R. Weir, 507 (in Mo. Bot. Gard. Herb., 

22700); Sidney, /. Macoun, 41 1 (in Mo. Bot. Gard. Herb., 
55311). 

Washington: Bingen, W. N. Suksdorf, 871; Falcon Valley, 

W. N. Suksdorf, 723; Hoquiom, C. J. Humphrey, 
Olympia, C. J. Humphrey, 6306; Renton, C. J. Humphrey 
6439. 

California: Muir Woods, W. A. Murrill, 1153 (in N. Y. Bot 

Gard. Herb, and Mo. Bot. Gard. Herb., 55705); Olema 
M. A. H. (in N. Y. Bot. Gard. Herb, aad Mo. Bot. Gard 
Herb., 56590); Sutro Woods, R. A. Harper (in N. Y. Bot 
Gard. Herb, and Mo. Bot. Gard. Herb., 56704). 

Arizona: Coronada Nat. Forest, Santa Catalina Mts., G. G 




[Vol. 7 



148 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Hedgcock & W. H. Long, comm. by C. G. Humphrey, 2561 
(in Mo. Bot. Gard. Herb., 9438). 



s. S. sulphuratum Berkeley & Ravenel, Linn. Soc. Bot. Jour 
331. 1868; Grevillea i: 163. 1873; Sacc. Syll 



566. 1888; Massee, Linn. Soc. Bot. Jour. 27 



1890. 
Plate 5, fig 



Stereurn ochroleucum Bresadola, Ann. Myc. 1 : 91 



1903 



Not 



Stereurn ochroleucum Fries, Hym 
ochroleucum Fries, Epicr. 557. 



1874. nor Corticium 



1838. 



Kew Herb, and Curtis Herb 



Fructifications coriaceous, stiff, effuso-reflexed, finally umbo 
,te alone the line of attachment to the substratum, and lobed 



upper 



surface toment 



con- 




Fig. 21. S. sulphuratum. 



:ally furrowed, "sulphur colored" 
when fresh, becoming cartridge-buff to 
gray in the herbarium, the surface not 
hardened and crust-like under the hairy 
covering; hymenium even, glabrous, 
becoming pinkish buff to dirty tilleul- 
buff in the herbarium; in structure 200- 
400 u thick under the hairy covering, 



Section of type x 68. The with the intermediate layer not differ 

outer border of intermediate entiated on itg upper side into a dens , 



layer not a colored, crust-like 



zone. 



golden zone but hyaline throu 



and with the 



tudinally 



ed 



hyphae 



diameter, curving outward on the upper side 



form the hirsute covering and curving downward on the under 



side to form the hymenium 



}red condu 



stidia ; spores hyal 



3 



M 



Fructifications with resupinate portion \-2 cm. broad, 10 cm 



more long on under side of limbs; reflexed lob 



cm 



broad 



2h cm 



On dead limbs of Betula and other frond 
Mexico. West Indies, Venezuela, and I 



Georgia 



September 



Jan 



Not common, 
wing condition, the 
S. sulvhuratum and 



pilei, taken in connection with geographic range wholly south 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 149 



of that of S. hirsutum, should render specimens of the former 
species easily distinguishable. All gatherings of S. sulphur atum 
which I have seen had already faded to the gray color of old, 
weathered S. hirsutum and in this condition are best distin- 
guished by not having underneath the hairy covering a thin 
hardened crust as the upper surface of the intermediate layer, 
nor a dense, somewhat golden zone on the upper border of the 
intermediate layer when sectional preparations are examined 
with the microscope. 

S. sulphuratum occurs also in Westphalia, Germany, appar- 
ently an isolated station, and has been confused there with 
Stereum ochroleucum Fries, a species of thicker and softer struc- 
ture having hyphae interwoven instead of densely and longitudi- 
nally arranged — for which reason Fries was doubtful about its 
being a true Stereum and published the species originally as a 
Corticium. Collections from Sweden and France communicated 
to me as cospecific with the Westphalian gatherings have the 
upper surface of the intermediate layers with a crust-like golden 
zone and are referable to S. hirsutum instead. 

Specimens examined: 
Exsiccati: Brinkmann, Westfalische Pilze, 49, under name of 

Stereum ochroleucum; Rick, Fungi Austro-Am., 260, under 

name of Stereum ochroleucum. 
Germany: Westphalia, Lengerich, W. Brinkmann, comm. by 

G. Bresadola, and in Brinkmann, Westfalische Pilze, 49. 
Georgia: Catoosa Springs, H. W. Ravenel (in Kew Herb, and in 

Curtis Herb., 1731). 
Florida: C.G.Lloyd, 2131. 

Alabama: Auburn, Ala. Biol. Surv., comm. by F. S. Earle; 

Montgomery, R. P. Burke, 4 (in Mo. Bot. Gard. Herb., 

22017). 
Mexico: Jalapa, W. A. & E. L. Murrill, 316, 343, comm. by 

N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 54438, 

55477). 

Cuba: C. Wright, 292, type (in Kew Herb.). 

Jamaica: Farr (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. 

Herb., 56667); Cinchona, W. A. & E. L. Murrill, 480, 
546, comm. by N. Y. Bot. Gard. Herb.; Morce's Gap, 
W. A. & E. L. Murrill, 723, comm. bv N. Y. Bot. Gard. 



150 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



Herb.; Monkey Hill, W. A. & E. L. Murrill, 784, comm. 
by N. Y. Bot. Gard. Herb.; Sir John Peak, L. M. Under- 
wood, 3182 (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. 
Herb., 56668). 

Venezuela: Fendler, 169 (in Curtis Herb.). 

Brazil: Sao Leopoldo, Rick, in Rick, Fungi Austro-Am., 260. 



S. hirsutum Willdenow ex Fries, Epicr. 5< 
39. 1874; Persoon, Roemer Neues M 



Eur. I 

1794; Obs. Myc. 2:90. 1799; Berkeley, Outlines 
270. pl.l7.f 



1838; Hym. 
Bot. i: 110. 



Sacc. Syll. Fung. 6 



5. 1888 
Plate 5. 



Thelephora hirsuta Willdenow, Fl. Berol. Prod. 397. 



Fries, Syst. Myc 
1801; Myc. Eur. 
Herb, de la France 



439 



1821 



Persoon, 




Fun 



1787 
570 



i: 116. 1822. — Auricularia refiexa Bulliard 



27A 



1785 



Schw 



Naturforscl 



G 



Thelephora ochracea 



but not of Fries 



sub 



Leipzig Schrift. i : 106. 1822, 



Elenchus 



1828; Schweinitz, Am. Phil. Soc. Trans. N. S 
Corticium subzonatum Fries, Epicr. 557. 1838 



1832. 



6: 

53 



1888. 
1914. 



Sacc. Syll 



— Stereum variicolor Lloyd, Myc. Writ. 4. Letter 



Berkeley, Outl. Brit. Fung. pi. 17. f. 7; Bolt 



Hist. Fung, pi 82; Bulliard, Herb, de la France, pi. 274; Hussey, 

111. Brit. Myc. 1 : pi. 58; Sowerby, 
Col. Figs. Brit. Fung, pi 27; 
Stevenson, Brit. Fungi 2: 




267 



/ 



See Sacc. Sy 



20: 890, for reference to other 
illustrations. 

Fructifications coriaceous, stiff, 



reflexed 



wholly 



hirsute 



some 



what concentrically furrowed 



complicate, cream-buff at first, 
Fig. 22. S. hirsutum. Section x becoming grayish when old and 

68; intermediate layer, * golden, wea thered, with a thin, hardened, 
crust-hke zone, 2; hymenmm contain- ... ' ' 

ing very few conducting organs, h; crust-like surface bearing the 
spores, s, x 488. hairy covering, the margin entire: 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII lOl 



hymenium even, warm buff at first, sometimes becoming pale 
smoke-gray, unchanged when cut or bruised; in structure 
500-700 n thick under the hairy covering, with the inter- 
mediate layer bordered next to the hairy covering by a very 
dense, narrow, golden zone, the rest of the intermediate layer 
composed of densely and longitudinally arranged hyaline hyphae 
3-4 m in diameter, some of which in the subhymenium are thick- 
walled, up to 5-6 n in diameter, and very rarely have golden- 
brown contents as seen between the basidia; no colored con- 
ducting organs, cystidia, nor gloeocystidia; spores white in spore 
collection, even, flattened on one side, 5-7|X2-S 

Reflexed portion varying from barely reflexed up to 2 cm. 
broad, 1-2 cm. long; fructifications merely gregarious or con- 
fluent, and imbricated. 

On logs and stumps of birch, beech, and other frondose 
species. Newfoundland to South Carolina and westward to 
British Columbia and California, and in Mexico. July to 
November in the east and to February in the Pacific states. 

Common. 
Stereum hirsutum is characterized by its strigose-hirsute, 

buff-colored pileus, weathering more or less gray, and by its 
warm buff hymenium, sometimes smoke-gray, which does not 
exude a red juice when wounded; as in S. rameale, S. versicolor, 
S. fasciatum, S. lobatum, S. australe, and S. gausapatum, the 
upper surface of the intermediate layer is differentiated into a 
thin, golden, somewhat horny crust from which the hairy cover- 
ing springs. This golden zone shows well under the microscope, 
and its presence is a decisive character for separating S. hir- 
sutum from the southern S. sulphur atum , a species of somewhat 

similar aspect. 

Specimens examined: 

Exsiccati: Berkeley, Brit. Fungi, 146; Ca.vara, Fungi Longo- 

bardiae, 61; Cooke, Fungi Brit., 108; Ellis, N. Am. Fungi, 
1204; Krieger, Fungi Sax., 118; Rabenhorst, Herb. Myc, 
211; Romell, Fungi Scand. Exs., 26. 

Sweden: Femsjo, L. Romell, two collections, and E. A. Burt; 

Mauritzberg, W. A. & E. L. Murrill, 4078 (in N. Y. Bot. 
Gard. Herb, and Mo. Bot. Gard. Herb., 56671); Stock- 
holm, L. Romell, 30, 401, and in Romell, Fungi Scand. 
Exs., 26. 



1 £0 {VoL - 7 

lOJ ANNALS OF THE MISSOURI BOTANICAL GARDEN 

England: M. J. Berkeley, in Berkeley, Brit. Fungi, 146; Epping, 

M. C. Cooke, in Cooke, Fungi Brit., 108; Kew Gardens, 

G. Massee; Selby, E. A. Burt. 
France: Fautrey, comm. by Lloyd Herb., 3326; Aveyron, A. 

Galzin, 8459, comm. by H. Bourdot, 7813; St. Priest, 

Allier, H. Bourdot, 19770. 

Germany: Nossen, Saxony, W . Krieger, in Krieger, Fungi Sax., 

118. 

Italy: A. Carestia, 784, 1215, comm. by G. Bresadola; Pavia, 

F. Cavara, in Cavara, Fungi Longobardiae, 61. 

Newfoundland: A. C. Waghorne, 118 (in Mo. Bot. Gard. Herb., 

5082). 

Canada: J. Macoun, 69. 

Ontario: Ottawa, J. Macoun, 16, 466a; Port Credit, /. H. Fault, 

Univ. Toronto Herb., 353 (in Mo. Bot. Gard. Herb., 44858) ; 

Toronto, G. H. Graham, Univ. Toronto Herb., 678 (in Mo. 

Bot. Gard. Herb., 44919). 
Maine: Milo, W. A. Murrill, 2024 (in N. Y. Bot. Gard. Herb. 

and Mo. Bot. Gard. Herb., 56682). 
New Hampshire: North Conway, L. 0. Overholts, 5009 (in Mo. 

Bot. Gard. Herb., 56346). 
Vermont: Middlebury, E. A. Burt; Ripton, E. A. Burt; Smug- 
glers Notch, E. A. Burt, two gatherings. 
Massachusetts: Boston, L. C. Monahan (in Mo. Bot. Gard. 

Herb., 15309); Cambridge, E. A. Burt; Mt. Auburn, 

E. A. Burt; Nahant, A. B. Seymour, T 81 (in Mo. Bot. 

Gard. Herb., 12954); Waverly, A. B. Seymour, T 25, T 26 

(in Mo. Bot. Gard. Herb., 16364, 18372); Waltham, A. B. 

Seymour, T 16 (in Mo. Bot. Gard. Herb., 17912). 
Connecticut: Broad Brook, C. C. Hanmer, 2682 (in Mo. Bot. 

Gard. Herb., 42606); Mansfield, P. W. Graff, 18 (in Mo. 

Bot. Gard. Herb., 44817); Storrs, P. W. Graff, 29 (in Mo. 



Gard. Herb 



New York 

by G. 



and W. H. Wright, comm. 
ove, C. L. Shear, 995: Fall 



Creek, W. H. Wright, 7992; Floodwood, E. A. Burt. 
Pennsylvania: Spruce Creek, J. H. Fault, Univ. Toronto Herb. 

337 (in Mo. Bot. Gard. Herb., 44883); West Chester 
Ever hart & Haines, in Ellis, N. Am. Fund. 1204. 



1920] 



BURT— THELEPHORACEAE OF NORTH AMERICA. XII 153 



North Carolina: Schweinitz, types of T. ochracea and T. sub- 

zonata (in Herb. Schweinitz) ; Blowing Rock, G. F. Atkinson 
4S08. 

South Carolina: Clemson College, P. H. Rolfs. 

Michigan: Cadillac, H. D. House, 1225 (in N. Y. Bot. Gard 

Herb, and Mo. Bot. Gard. Herb., 56673); Isle Royale 
Miss A. D. Stucki, Univ. Wis. Herb., 23; Vermilion, A. H 
W. Povah, 199 (in Mo. Bot. Gard. Herb., 15145). 

Indiana : Crawfordsville, D. Reddick, 5, 7, and another specimen 

comm. by H. H. Whetzel. 

West Virginia: Paw Paw, C. L. Shear, 1173. 

Tennessee: Elkmont, C. H. Kaufman, 62 (in Mo. Bot. Gard. 

Herb., 3972). 

Wisconsin: Blue Mounds, Miss A. D. Stucki, Univ. Wis. Herb., 

8, 9; Madison, Miss A. D. Stucki, Univ. Wis. Herb., 34, 
and W. Trelease, 5, 26 (in Mo. Bot. Gard. Herb., 56683, 
56684); Palmyra, Miss A. D. Stucki, Univ. Wis. Herb., 33. 

Minnesota: Lake Itaska, comm. by E. L. Jensen, 9 (in Mo. Bot. 

Gard. Herb., 11088). 

Missouri: B. M. Duggar, 95; Meramec, P. Spaulding (in Mo. 

Bot. Gard. Herb., 5025). 

Arkansas: Womble, W. H. Long, 198U, 19883 (in Mo. Bot. 

Gard. Herb., 8963, 14651). 
Nebraska: Lincoln, C. L. Shear, 1023. 
Montana: Evaro, J. R. Weir, 431 (in Mo. Bot. Gard. Herb., 

22515); Mystic Lake, C. L. Shear, 1102. 
Colorado: Steamboat Springs, E. Bartholomew, 5578 (in Mo. 

Bot. Gard. Herb., 9185, 44584); Tolland, F. J. Seaver & 

E. Bethel (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. 

Herb., 56674). 

New Mexico: Albuquerque, W. H. Long, 21153 (in Mo. Bot. 

Gard. Herb., 55112); Cloudcroft, F. S. Earle, 495, comm. 
by N. Y. Bot. Gard. Herb., and W. H. Long, 19542 (in 
Mo. Bot. Gard. Herb., 55111); Tejano Exp. Station, W. 
H. Long, 21875, 21894, 21907 (in Mo. Bot. Gard. Herb., 
55161-55163); Tyom Exp. Station, W. H. Long, 21365, 
21366, 21426 (in Mo. Bot. Gard. Herb., 55113, 55114, 
55160); Ute Park, P. C. Standley, 14197, comm. by N. Y. 
Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 44953) ; Weeds, 



[Vol. 7 



154 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



L. Wymans, comm. by W. H. Long, 12969 (in Mo. Bot. 

Gard. Herb., 55110). 

Idaho: Priest River, J. R. Weir, 19,31, 48. 

British Columbia: New Westminster, A. I. Hill (in N. Y. Bot. 

Gard. Herb, and Mo. Bot. Gard. Herb., 56675) ; Oak Bay, 
/. Macoun, 579a (in Mo. Bot. Gard. Herb., 55310) ; Sidney, 
/. Macoun, 46, 47, 49, 52, 52 bis, 53, 54, 84 (in Mo. Bot. 
Gard. Herb., 5736, 6674, 6694, 6682, 55361, 6698, 6697, 

6704 respectively). 

Washington: Bingen, W. N. Suksdorf, 692, 693, 709, 874, 891, 

893, 916, 953; Kalama, C. J. Humphrey, 61 40; Chehalis, 
C. J. Humphrey, 6254 (in Mo. Bot. Gard. Herb., 16677); 
Olympia, C. J. Humphrey, 6310; Seattle, S. M. Zeller, 119 
(in Mo. Bot. Gard. Herb., 44139); Tacoma, W. A. Murrill, 
127, 142, comm. by N. Y. Bot. Gard. Herb, (in Mo. Bot. 
Gard. Herb., 55744, 55730). 

Oregon: Corvallis, C. E. Owens, 2036, 2054, 2057, 2084, 2135, 

2136, 2139, 2142, 2143 (in Mo. Bot. Gard. Herb., 43872, 
43878, 43877, 44249, 44695, 44694, 44693, 44699, 44702 
respectively), and S. M. Zeller, 181 4 (in Mo. Bot. Gard. 
Herb., 56332); Eugene, C. J. Humphrey, 6050, 6063, 6076 
(in Mo. Bot. Gard. Herb., 17175); Mt. Hood, G. G. Hedg- 
cock, comm. by C. J. Humphrey, 2569 (in Mo. Bot. Gard. 
Herb., 16418); Granite Pass, J. R. Weir, 8680, 8681 (in 
Mo. Bot. Gard. Herb., 36752, 36753). 

California: R. A. Harper, 8, 109, 141, 148 (in N. Y. Bot, Gard. 

Herb, and Mo. Bot. Gard. Herb., 56678-56681), and Miss 
E. Hyatt, comm. by C. L. Shear, 1089; Berkeley, C. J. 
Humphrey, 5970, 5982, H. A. Lee, Univ. Calif. Herb., 1015, 
1016, 1019, 1021, 1022 (in Mo. Bot. Gard. Herb., 44154- 
44156, 44152, 44157 respectively), W. A. Setchell, Univ. 
Calif. Herb., 1023, 1024 (in Mo. Bot. Gard. Herb., 44153, 
44245), and G. Courvoisier, Univ. Calif. Herb., 1025 (in Mo. 
Bot. Gard. Herb., 44149) ; Claremont, D. L. Crawford, D 12, 
comm. by L. O. Overholts, 3280 (in Mo. Bot. Gard. Herb., 
10479) ; Coast Range, C. F. Baker, 82, 101, comm. by N. Y. 
Bot. Gard. Herb.; Fair Oaks, R. A. Harper (in N. Y. Bot. 
Gard. Herb, and Mo. Bot. Gard. Herb., 56676); Julian, 
E. Bethel. 28272 (in Mo. Bot. Gard. Herb.. 55439): North- 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



155 



brae, L. S. Smith, Univ. Calif. Herb., 416 (in Mo. Bot. Gard. 
Herb., 44148); Muir Woods, W. A. Murrill, 1133 (in N. Y. 
Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 55713); Pine- 
hurst, E. Bethel, 26269, 26274 (in Mo. Bot. Gard. Herb., 
55438, 55440); Preston's Ravine, W. A. Murrill & L. S. 
Abrams, 1171, comm. by N. Y. Bot. Gard. Herb, (in Mo. 
Bot. Gard. Herb., 55707); San Francisco, W. A. Setchell & 
C. C. Dolier, W. A. Murrill, 1111, comm. by N. Y. Bot. 
Gard. Herb, (in Mo. Bot. Gard. Herb., 55702); Santa 



Barbara, 0. M. Oleso 



Santa Cruz, G. J. Str eater 



N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb 



56677); Sutro Forest, A. S. Rhoads, 1 (ir 
Herb., 56045). 
Mexico: Coyoacan, Roldan, comm. by J. R. "W 

(in Mo. Bot. Gard. Herb.. 56795. 56796). 



Mo. Bot. Gard 



44. S. fasciatum Schweinitz, Naturforsch. Ges. Leipzig 
Schrift. 1: 106. 1832 (under B. Sterea of Thelephora) ; Fries, 
Epicr. 546. 1838 Sacc. Syll. Fung. 6: 560. 1888; Massee, 
Linn. Soc. Bot. Jour. 27: 180. 1890. 

Plate 5, figs. 43-45. 
Thelephora versicolor fasciata (Schw.) Fries, Elenchus Fung. 
1: 175. 1828; Schweinitz, Am. Phil. Soc. Trans. N. S. 4: 167. 
1832. — T. ostrea Blume & Nees, Acad. Leop.-Carol. Nov. Acta 
I3 1 : 13. pi 2. 1S2Q.— Stereum ostrea (Bl. & Nees) Fries, 
Epicr. 547. 1838; Sacc. Syll. Fung. 6:571. 1888; Bresa- 
dola, Hedwigia 51: 321. 1912. — Thelephora (Stereum) mollis 
Leveille, Ann. Sci. Nat. Bot. III. 5: 147. 1846. — Stereum 
molle Leveille in Sacc. Syll. Fung. 6: 577. 1888; Massee, 
Linn. Soc. Bot. Jour. 27: 175. 1890 — 
arcticum Fries, Hym. Eur. 639. 1874. 



o. 



Type: in Herb. Schweinitz and in Curtis 
Herb. 

Fructifications coriaceous, rigid, in the 
north at first broadly effuso-reflexed with 
the resupinate portion narrow, soon umbo- 
nate sessile — perhaps so from the first 
in the tropics — often laterally confluent, 




sometimes pseudo-stipitate by prolongation x 665. 



Fig. 23. S. fasciatum. 
Section of reflexed stage, 
natural size; spores, s, 



[Vol. 7 
156 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



of the umbo, at first densely tomentose and drying warm buff to 
tawny olive, at length weathering to pale smoke-gray to neutral 
gray and sometimes with the tomentum torn apart in narrow 
zones and showing the hazel or chestnut surface of the bared 
areas, the margin normally entire; in structure 400-700 m thick, 
with the intermediate layer composed of very densely arranged, 
hyaline hyphae 4 n in diameter and bordered on the upper side 
by a broad dark zone which bears the tomentum of the upper 
surface; hymenium glabrous, usually warm buff to cinnamon- 
buff, sometimes assuming violaceous tints; no cystidia, gloeo- 
cystidia, nor conducting organs; spores from spore collections 
white, even, flattened on one side, 5^-7§X2|-3 n. 

Fructifications 2-7 cm. in diameter, often laterally confluent. 

On logs and stumps of Quercus and other hardwood species. 
Common throughout North America from Canada southward, 
in the West Indies, and in South America; occurs also in Norway, 
Sweden, Formosa, and Java, although apparently rare in the 
Old World. In vegetative condition from June onward in the 
north, persisting throughout the year. 

Specimens of S. fasciatum may be distinguished from those of 
the less common S. lobatum by the thicker tomentose covering 
of the former, which may continue unbroken throughout the 
year or become torn apart so as to show rather few and narrow, 
bared chestnut zones; the pileus of S. fasciatum is thicker than 
that of S. lobatum, and the margin has a lobate tendency but 
rarely. Towards the northern part of its range where I have 
observed the development of fructifications throughout the 
season, the fructifications are at first effuso-reflexed with the 
resupinate portion up to 1 cm. broad, the reflexed portion 1^ 
cm. from base to margin, and with a lateral extent along the 
substratum of 2-8 cm.; umbos soon form at points 1-2 cm. 
apart along line of intersection of the plane of reflexed portion 
with the substratum; by further growth outward of the laterally 
confluent pilei these umbos become the final points of attach- 
ment of the pilei with the substratum. In Washington and 
California the fructifications may continue broadly reflexed 
when old and are difficult to distinguish from luxuriantly grown 
S. hirsutum. 

The specimens from Formosa, cited below, are in the stage in 






1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 157 



which the fructifications are still with a resupinate portion but 
with the umbos distinctly outlined, and exactly agree in all 
respects, even including spore dimensions, with my Vermont 
collections of the same stage. The authentic specimen of 
Thelephora ostrea from Java is in the final stage with attachment 
by umbo only and is clothed over its whole upper surface with a 
thick coat of tomentum, and matches well most of the specimens 
of the type collection of Stereum fasciatum in Herb. Schweinitz. 
I infer from the lack of specimens of S. fasciatum from the East 
Indies and the Philippines in published exsiccati, that this 
species is very rare there and that what frequently has been 
listed as S. ostrea is really the very common S. concolor instead. 
Schweinitz's original description of S. fasciatum presents at 
such length the disappearance of tomentum from the upper 
surface of the pileus and the broad, glabrous, shining surface 
with many vari-colored zones, that it seems probable he may 
have intended the description to comprehend not only S. fascia- 
tum as treated by me but also S. lobatum, which he must have 
seen about him in North Carolina; nevertheless, the ample 
collection of specimens in Herb. Schweinitz which were preserved 
as the type of S. fasciatum contains no fructifications referable 
to S. lobatum. 

Specimens examined: 
Exsiccati: Bartholomew, Fungi Col., 2590, under the name S. 

versicolor, 2884, under the name S. versicolor, 2985, 3985, 
4291, and 4986; Ellis, N. Am. Fungi, 18, under the name 
S. versicolor v. fasciata, 514a, and c, both under the name 
S. versicolor; Ell. & Ev., N. Am. Fungi, 1714, under the 
name S. purpureum; Ellis & Ev., Fungi Col., 306, under the 
name S. versicolor; Ravenel, Fungi Am., 220, under the 
name S. versicolor, and 721; Smith, Central Am. Fungi, 
145, under the name S. versicolor; de Thiimen, Myc. Univ., 
2011, mixed with S. lobatum. 
Norway: Bosekon, Finmark, M. N. Blytt, type of Stereum 

arcticum (in Herb. Fries). 
Sweden: on Alnus, North Sweden, comm. by L. Romell, 400. 
Canada: J. Macoun, 12. 

Prince Edward Island: J. Macoun, 846 (in Macoun Herb.). 
Quebec: J. Macoun, 77, 239, 249, 464 (all in Macoun Herb.) 



1 ec [VoL - 7 

105 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Ontario: Bond Lake, J. H. Faull, Univ. Toronto Herb., 319 

(in Mo. Bot. Gard. Herb., 44874); Ottawa, J. Macoun, 50; 
Port Credit, J. H. Faull, Univ. Toronto Herb., 352, 354 
(in Mo. Bot. Gard. Herb., 44857, 44856); Rondeau Park, 
J. H. Faull, Univ. Toronto Herb., 358 (in Mo. Bot. Gard. 
Herb., 44870); Toronto, J. H. Faull, Univ. Toronto Herb., 
356 (in Mo. Bot. Gard. Herb., 44868), T. Langton, Univ. 
Toronto Herb., 501 (in Mo. Bot. Gard. Herb., 44853), G. H. 
Graham, Univ. Toronto Herb., 680 (in Mo. Bot. Gard. 
Herb., 44937). 

Maine: Harrison, J. Blake, comm. by P. L. Ricker; Orono, 

F. L. Harvey, comm. by P. L. Ricker; Portage, L. W. 

Riddle, 2,17. 

Vermont: Middlebury, E. Brainerd, E. A. Burt, nine collections; 

Ripton, E. A. Burt. 

Massachusetts: Amherst, P. J. Anderson, 2, 4 (in Mo. Bot. 

Gard. Herb., 55846, 55845 respectively). 
Connecticut: Mansfield, P. W. Graff, 30 (in Mo. Bot. Gard. 

Herb., 44803); New Haven, W. A. Setchell; Norwich, W. 

A. Setchell. 

New York: Sartwell, 19 (in Mo. Bot. Gard. Herb., 5076); 

Alcove, C. L. Shear, 1327; Canandaigua, L. M. Under- 
wood, 21, distributed under the name S. versicolor (in Mo. 
Bot. Gard. Herb., 5117); East Galway, E. A. Burt; Flood- 
wood, E. A. Burt; Freeville, G. F. Atkinson, 2821; Glasco, 
P. Wilson, 48, 43 (in Mo. Bot. Gard. Herb., 54752, 54754); 
Grand View, H. vonSchrenk (in Mo. Bot. Gard. Herb.. 42811. 



G. F. Atkinson, 2819, 2820, 8027, Bot. 



Dept. Cornell Univ., 133 0. S., 2871, H. S. Jackson, comm. 

by Bot. Dept. Cornell Univ., 14397-14399, Van Hook, 
comm. by Bot. Dept. Cornell Univ., 8084, W. C. Muenscher, 
147, 205, 211 (in Mo. Bot. Gard. Herb., 56602-56604); 
Palisades, P. Wilson, 20, 18, 12 (in Mo. Bot. Gard. Herb.. 



54759) ; Yonkers, P. Wilson, 61 (in M 



Gard. Herb 



New Jersey: Alpine, P. Wilson, 17, 13, 7 (in Mo. Bot. Gard. 

Herb., 54757, 54758, and 54760 respectively); Belleplain, 
C. L. Shear, 1250; Newfield, J. B. Ellis, in Ellis, N. Am. 
Fungi, 18, 514c, and Ell. & Ev., Fungi Col., 306. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 159 



Pennsylvania: E. Michener, 88 (in Mo. Bot. Gard. Herb., 5044); 

Germantown, E. A. Burt; Huntington Co., A. S. Rhoads, 7 
(in Mo. Bot. Gard. Herb., 44084); Lancaster City, Mrs. 
A. F. Eby (in Mo. Bot. Gard. Herb., 5083); Kittanning, 
D. R. Sumstine, 4, 7, 7', Philadelphia, A. S. Rhoads, 19 
(in Mo. Bot. Gard. Herb., 44096) ; in coal mine, Pottsville, 
C. J. Humphrey, 310; Spruce Creek, J. H. Faull, Univ. 
Toronto Herb., 357, 359, 334, 670, 355, 667 (in Mo. Bot. 
Gard. Herb., 44869, 44871, 44888, 44917, 44926, and 44934 
respectively); Shingleton Gap, A. S. Rhoads, 15 (in Mo. 
Bot. Gard. Herb., 44093); State College, C. R. Orton, 1, 18 
(in Mo. Bot. Gard. Herb., 44079, 44095), comm. by L. O. 
Overholts, 2658, 5003 (in Mo. Bot. Gard. Herb., 5721, 
56345), A. S. Rhoads, 16 (in Mo. Bot. Gard. Herb., 44094); 
Trexlertown, C. G. Lloyd, 0084; in coal mine, Wadesville 
Colliery, C. J. Humphrey, 21583. 

Maryland: Glen Sligo, C. L. Shear, 1133. 

District of Columbia: Takoma Park, P. L. Richer, 820, C. L. 

Shear, 956. 

Virginia: Great Falls, 0. F. Cook, comm. by P. L. Ricker; Mt. 

Vernon, P. L. Ricker, 1121 in part; Mountain Lake, W. A. 

Murrill, 408 (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. 

Herb., 56618); Norton, A. B. Seymour (in Mo. Bot. Gard. 

Herb., 16405). 
North Carolina: Schweinitz, type (in Herb. Schweinitz and 

Curtis Herb.); Blowing Rock, G. F. Atkinson, 41^8, 4180, 

4315; Chapel Hill, W. C. Coker, 938 (in Mo. Bot. Gard. 
Herb., 56665); Leicester, B. B. Higgins, in Bartholomew, 
Fungi Col., 2985. 
South Carolina: Clemson College, P. H. Rolfs, 1613, 1616, 1619, 

1620, 1624, 1629, 1631, 1635. 

Georgia: Darien, H. W. Ravenel, in Ravenel, Fungi Am., 220, 

721; Dixie, R. M. Harper, 1633b, comm. by N. Y. Bot. 
Gard. Herb.; Tallulah Falls, A. B. Seymour, comm. by 

W. G. Farlow, 6 (in Mo. Bot. Gard. Herb., 55290). 
Florida: C. G. Lloyd (in Mo. Bot. Gard. Herb., 44068); Cocoa- 
nut Grove, H. von Schrenk (in Mo. Bot. Gard. Herb., 
43097); Eustis, L. M. Underwood, 1368, 1801 (in N. Y. 
Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 56616, 56617). 






(Vol. 7 
160 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Alabama: Adger, C. J. Humphrey; Montgomery Co., R. P. 

Burke, 34 (in Mo. Bot. Gard. Herb., 4273); Maplesville, 

C. S. Hill, comm. by C. J. Humphrey, 251. 
Mississippi: Laurel, C. J. Humphrey, 5431,5435; Ocean Springs, 

F. S. Earle (in Mo. Bot. Gard. Herb., 5118). 
Louisiana: Baton Rouge, C. W. Edgerton, 848, comm. by C. J. 

Humphrey; St. Martinville, A. B. Langlois, 2902, bf. 
Ohio: Cincinnati, D. L. James, in Ellis, N. Am. Fungi, 514c, 

C. G. Lloyd, 1579, 4499, 4501, 4506; Columbus, W. A. 

Kellerman, in Kellerman, Ohio Fungi, 33, under the name 

- 

S. versicolor; Granville, H. L. Jones; Linwood, C. G. Lloyd, 
2436, 02821, 02830; Penfield, F. D. Kelsey (in Mo. Bot. 
Gard. Herb., 5075); Worthington, Dr. Paddock (in Mo. 
Bot. Gard. Herb., 5114, 5157). 
Kentucky: Bowling Green, Miss S. F. Price (in Mo. Bot. Gard. 



Herb 



Mammoth Cave, C. G. Lloyd 



Tennessee: Algood, C. J. Humphrey, 308. 

Michigan: Isle Royale, Allen & Stuntz, 22, 60; Sailor's Encamp- 
ment, E. T. & S. A. Harper, 710; Vermilion, A. H. W. 
Povah, 142 (in Mo. Bot. Gard. Herb., 15144). 

Wisconsin: Bayfield, V. B. Walker, 6b (in Mo. Bot. Gard. Herb., 

9733); Blanchardville, Miss A. 0. Stucki, 47; Blue Mounds, 
Miss A. 0. Stucki, 49; Ithaca, W. Trelease, 89 (in Mo. 
Bot. Gard. Herb., 56606); Madison, E. T. Bartholomew, in 
Bartholomew, Fungi Col., 3985, Miss A. 0. Stucki, 31, 35, 
36, 50, W. Trelease (in Mo. Bot. Gard. Herb., 56605) ; Syene, 
W. Trelease, 90 (in Mo. Bot. Gard. Herb., 5072). 

Indiana: Greencastle, L. M. Underwood, 2 (in Mo. Bot. Gard. 

Herb., 44101); Hibernian Mills, Whetzel & Reddick, comm. 
by D. Reddick, 6, 8; Ladoga, P. J. Anderson, 1 (in Mo. 
Bot. Gard. Herb., 55838); Wabash "bottom", W. Trelease 
(in Mo. Bot. Gard. Herb., 5073). 

Illinois: Brownsville, E. T. & S. A. Harper, 951; Cobden (in 

Mo. Bot. Gard. Herb., 44102); Grand Pass Club, W. 
Trelease (in Mo. Bot. Gard. Herb., 5053); Jacksonville, 
E. Bartholomew, in Bartholomew, Fungi Col., 2590. 

Missouri: Bismarck, L. O. Overholts (in Mo. Bot. Gard. Herb.. 



43702) ; 



Gard. Herb 



5131); Columbia, B. M. Duggar, 346a, 562, 580; Creve 



« 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 161 



Coeur, E. A. Burt (in Mo. Bot. Gard. Herb., 8727) ; Lincoln 

Co., C. Trenning (in Mo. Bot. Gard. Herb., 4098); Mera- 

mec, P. Spaulding, 1, and (in Mo. Bot. Gard. Herb., 5020), 

Spaulding & Johnson (in Mo. Bot. Gard. Herb., 5013-5015); 

Meramec Highlands, N. M. Glatfelter (in Mo. Bot. Gard. 

Herb., 42583) ; Old Orchard, L. H. Pammel (in Mo. Bot. 

Gard. Herb., 5020, 5041); Piedmont (in Mo. Bot. Gard. 

Herb., 4783); Upper Creve Coeur, E. A. Burt (in Mo. Bot. 

Gard. Herb., 44057); Valley Park, H. von Schrenk (in Mo. 

Bot. Gard. Herb., 42859); White House, E. A. Burt (in 

Mo. Bot. Gard. Herb., 43808), contains mesopod specimen; 

Willow Springs, H. von Schrenk, 1, 2 (in Burt Herb, and 

Mo. Bot. Gard. Herb., 42886, 42851). 
Arkansas: Bertig, W. Trelease (in Mo. Bot. Gard. Herb., 5148); 

Big Flat, W. H. Long, 19859 (in Mo. Bot. Gard. Herb., 

8268); Fayetteville, E. Bartholomev), in Bartholomew, 

Fungi Col., 2884; Womble, W. H. Long, 19866 (in Mo. 

Bot. Gard. Herb., 8889); Wynne, W. Trelease (in Mo. 

Bot. Gard. Herb., 5147, 5152). 
Oklahoma: Poteau, W. Trelease (in Mo. Bot. Gard. Herb., 

5052) ; Spiro, E. Bartholomew, in Bartholomew, Fungi Col., 

4291. 

Texas: L. H. Pammel (in Mo. Bot. Gard. Herb., 56607) ; Austin, 

W. H. Long, Jr., 739; Gillespie County, G. Jermy (in Mo. 
Bot. Gard. Herb., 5048-5050) and 4^, comm. by U. S. 
Dept. Agr. Herb.; Joaquin, E. Bartholomew, in Bartholo- 
mew, Fungi Col., 4986; Quitman, W. H. Long, 12099 (in 
Mo. Bot, Gard. Herb., 55126) ; Waco, W. H. Long, Jr., 508. 

South Dakota: Black Hills, J. R. Weir, 10012 (in Mo. Bot. 

Gard. Herb., 55793). 

Nebraska: Memphis, T. A. Williams, comm. by C. L. Shear, 

1059; Nebraska City, V. B. Walker, 10 (in Mo. Bot. Gard. 

Herb., 12963). 
Kansas: Bourbon County, A. G. Barrett, 112, 115, 126, 127; 

Topeka, E. T. & S. A. Harper, 753. 

Colorado: Golden, Bethel & Overholts, comm. by L. O. Over- 
holts, 1758 (in Mo. Bot. Gard. Herb., 54871). 

New Mexico: Cloudcroft, F. S. Earle, 495 (inN. Y. Bot. Gard. 



Herb, and Mo. Bot. Gard. Herb., 1540). 



6 



[Vol. 7 
162 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Montana: Moeville, J. A. Hughes, comm. by J. R. Weir, 9750 

(in Mo. Bot. Gard. Herb., 56225). 
Idaho: Moscow, J. R. Weir, 7946 (in Mo. Bot. Gard. Herb., 

56218); Priest River, J. R. Weir, 6, 11, 49. 
British Columbia: Secamons, J. Macoun, 166; Sidney, J. Macoun, 

57, 70, 71 (in Mo. Bot. Gard. Herb., 5739, 5746, 5747). 
Washington: Bingen, W. N. Suksdorf, 694; Friday Harbor, 

V. B. Walker, 2 (in Mo. Bot. Gard. Herb., 8359); Lake 

Waldemen, C. H. Kauffman (in Mo. Bot. Gard. Herb., 

20763); Seattle, S. M. Zeller, 68, 118 (in Mo. Bot. Gard. 

Herb., 44137, 44143); Tacoma, E. Bartholomew, 4929 (in 

Mo. Bot. Gard. Herb., 20810). 
Oregon: Corvallis, C. E. Owens, ~„„~. 





, 2055, 2140, 2141 
(in Mo. Bot. Gard. Herb., 43874-43876, 44700, 44701); 
Granite Pass, J. R. Weir, 8675 (in Mo. Bot. Gard. Herb., 
36750); Wallowa, C. J. Humphrey, 265; Siskiyou National 
Forest, J. R. Weir, 8678 (in Mo. Bot. Gard. Herb., 36751). 

California: R. A. Harper, 39, 108, 142 (in N. Y. Bot. Gard. 

Herb, and Mo. Bot. Gard. Herb., 56610-12); C. R. Orcutt, 
in Ell. & Ev., N. Am. Fungi, 714; La Honda, Edna Hyatt, 
comm. by C. L. Shear, 1088, 1091; Muir Woods, W. A. 
Murrill, 1158, comm. by N. Y. Bot. Gard. Herb, (in Mo. 
Bot. Gard. Herb., 55715); Redding, C. J. Humphrey, 1035; 
San Francisco, A. S. Rhoads, 2 (in Mo. Bot. Gard. Herb., 
56046); Saratoga, E. B. Copeland, 1806. 

Arizona: Crown King, G. G. Hedgcock, comm. by C. J. Hum- 
phrey, 2564 (in Mo. Bot. Gard. Herb., 10752). 

Mexico: Cordoba, W. A. & E. L. Murrill, 996, comm. by N. Y. 

Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 54609) ; Guerna- 
vaca, W. A. & E. L. Murrill, 415, 416, 412, comm. by N. Y. 
Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 54518, 54519, 
54543); Jalapa, W. A. & E. L. Murrill, 75, 148, 193, 
comm. by N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 
11275, 10360, 54436), C. L. Smith, in Smith, Central Am. 
Fungi, 145; Oaxaca, E. W. D. Holway; Orizaba, W. A. & 
E. L. Murrill, 758, comm. by N. Y. Bot. Gard. Herb, (in 
Mo. Bot. Gard. Herb., 54632); Parral, E. O. Matthews (in 
Mo. Bot. Gard. Herb., 5722, 10459). 

Guatemala: Maxon & Hay, 3250, comm. by U. S. Bur. PI. Ind. 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 163 



Honduras: P. Wilson, 188, comm. by N. Y. Bot. Gard. Herb. 
Cuba: Ciego de Avila, Earle & Murrill, 568, comm. by N. Y. 

Bot. Gard. Herb.; Fecha, F. S. Earle, 146, Earle & Wilson, 

224; Guantanamo, J. R. Weir, 10644 (in Mo. Bot. Gard. 

Herb., 56237); Oriente, J. A. Shafer, 3392, 8468 (in N. Y. 

Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 56613, 56614); 

San Diego de los Baiios, Earle & Murrill, 331, comm. by 

N. Y. Bot. Gard. Herb. 

Porto Rico: Bayamon, /. A. Stevenson, 5427 (in Mo. Bot. Gard. 

Herb., 8180); Mayaguez, F. S. Earle, 89, comm. by N. Y. 

Bot. Gard. Herb.; Rio Piedras, Johnston & Stevenson, 

comm. by J. A. Stevenson, 1764, 1937, 2005 (in Mo. Bot. 

Gard. Herb., 9824, 14220, 14270); San Jaun, Mr. & Mrs. 

A. S. Heller, 700, comm. by N. Y. Bot. Gard. Herb. 
Jamaica: Cinchona, W. A.& E.L. Murrill, 450, 499, 521, comm. 

by N. Y. Bot. Gard. Herb., H. von Schrenk (in Mo. Bot. 

Gard. Herb., 43630); Chester Vale, W. A. & E. L. Murrill, 

282, 316, comm. by N. Y. Bot. Gard. Herb.; Monkey Hill, 

W. A. Murrill, 817, comm. by N. Y. Bot. Gard. Herb.; 

Moore Town, W. A. & E. L. Murrill, 160, comm. by N. Y. 

Bot. Gard. Herb. 
Brazil: Malme (in Romell Herb.). 

Formosa: Urai, S. Kusano, 11.16 (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56587). 
Java: Junghuhn, authentic specimen of Thelephora ostrea, comm. 

by G. Bresadola. 

Philippine Islands: Luzon, H. M. Curran, Forestry Bureau, 

9665 (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 
56583); Mindanao, A. D. E. Elmer, 10556, Philippine Is. 
PI. (in Mo. Bot. Gard. Herb., 705743). 



45. S. lobatum (Kunze) Fries, Epicr. 547. 1838; Sacc. Syll. 
Fung. 6: 568. 1888; Massee, Linn. Soc. Bot. Jour. 27: 175. 

Plate 5, fig. 46. 



1890. 

Thelephora lobata Kunze in Weigelt Exsiccati, 1827; Fries, 
Linnaea 5: 527. 1830. — Stereum Sprucei Berk. & Curtis, Linn. 
Soc. Bot. Jour. 10: 331. 1868; Sacc. Syll. Fung. 6: 567. 1888. 
— An S. concolor Junghuhn, Crypt. Java, 38. 1838? See Sacc. 
Syll. Fung. 6: 561. 1888; Bresadola, Hedwigia 51 : 321. 1912. 



[Vol. 7 



164 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 




Illustrations: Engl. & Prantl, Nat. Pflanzenfam. (i: 1** 
124. text /. 69, A-B; Hard, Mushrooms, 455. text f. 382, 
as S. versicolor. 

Type: type distribution in Weigelt Exs. 

Fructifications coriaceous, rigid, thin, wedge-shaped to um- 
bonate, sessile, often laterally concrescent, at first tomentose 
and drying tawny olive, at length with the tomentum becoming 
pale smoke-gray to whitish, disappearing more or less near the 
margin and in narrow zones and showing the glabrous, shining, 
hazel surface of the bared areas, the margin undulate and 
usually more or less lobed; in structure 300 ju thick, with the 
intermediate layer composed of densely arranged, thick-walled r 
hyaline hyphae 4-4-2- m in diameter; hymenium glabrous, even, 
usually drying pinkish buff; no setae, gloeocystidia, nor con- 
ducting organs; spores hyaline, even, flattened on one side, 

4-5X1 ^-2 m, but few seen. 

Pileus usually 3-7 cm. long, 2-6 cm. broad, sometimes much 

larger by lateral confluence. 

On dead branches, logs, and stumps of frondose species in 
the cases noted. A tropical species ranging northward to New 
York and Wisconsin and southward to Brazil. Occurs in the 
Philippine Islands and East Indies also, if S. concolor is a synonym. 

S. lobatum may be distinguished from the related S.fasciatum, 
S. versicolor, and S. radians by having a more or less lobate 
pileus which is also very thin, somewhat flexible, zonate on the 
upper side, with glabrous, shining hazel zones alternating with 
whitish tomentose zones of soft, matted hairs. No specimens 
of this species which I have examined have the pileus effuso- 
reflexed when young. Specimens of S. fasciatum occasionally 
have a somewhat lobate margin but the pileus is thicker, more 
heavily clothed with a tomentum which is more persistent than 
that of S. lobatum, and in its more northern stations where I 
have been able to observe the development, the young fructifica- 
tions are often effuso-reflexed at first. 

S. lobatum is primarily an American species described from 
collections made in Surinam, Dutch Guiana, but it seems prob- 
able that this species has a more extended geographical range 
through the tropical lands of the Eastern Hemisphere also. 
The recent collections in Philippine Islands, determined by 



1920) 

BURT — THELEPHORACEAE OF NORTH A.MERICA. XII 165 



Bresadola as S. concolor (Jungh.) and distributed in Elmer, 
Philippine Islands Plants, show that this species is but slightly, 
if at all, different from S. lobatum. The general aspect is the 
same but the Philippine specimens are the larger; none of them 
have their tomentum as soft and whitish as in S. lobatum. 
Some of these specimens have shown in crushed preparations 
spore-like bodies 3 m in diameter; spore collections of oriental 
specimens should be made. 

Specimens examined: 
Exsiccati: Bartholomew, Fungi Col., 4586, under the name S. 

fasciatum; Ellis, N. Am. Fungi, 514b, under the name 

S. versicolor v. fasciata, 514d, under the name S. versicolor v. 

petaliforme ; Ravenel, Fungi Car. i: 28, mixed with S. 

fasciatum; de Thiimen, Myc. Univ., 2011, mixed with S. 

fasciatum. 
New York: Alcove, C. L. Shear, 1019; Ithaca, L. A. Zinn, 82a 

(in Mo. Bot. Gard. Herb., 43074). 
Pennsylvania: West Chester, /. B. Gray, in Ellis, N. Am. Fungi, 

514b. 
North Carolina: Black Oak, H. W. Ravenel, in Ravenel, Fungi 

Car. i: 28; Blowing Rock, G. F. Atkinson, 4811, 4314; 
Chapel Hill, W. C. Coker, 331 (in Mo. Bot. Gard. Herb., 
56663); Transylvania County, W. A. Murrill & H. D. 
House, 425 (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. 

Herb., 56602). 

Georgia: Flint River, R. M. Harper, 1401a comm. by N. Y. Bot. 

Gard. Herb, (also in Mo. Bot. Gard. Herb., 5087); Dixie, 
R. M. Harper, 1633 (in Mo. Bot. Gard. Herb., 56603). 

Florida: C. G. Lloyd, 4833; Crescent City, Dr. G. Martin, in 

Ellis, N. Am. Fungi, 514d; Eustis, G. V. Nash, 2128 (in 
Mo. Bot. Gard. Herb., 5118), and L. M. Underwood, 1371 
(in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 
56601); Lake City, P. L. Ricker, 893; New Smyrna, C. G. 
Lloyd, 183; Tallahassee, E. Bartholomew, in Bartholomew, 

Fungi Col., 4586. 
Alabama: Auburn, F. S. Earle, from Lloyd Herb., 3459; Che- 
haw, E. A. Burt, two collections; Fayette Co., P. V. Sig- 
gers, comm. by A. H. W. Povah, 14 (in Mo. Bot. Gard. 
Herb., 9229). 



i aa rv° L - 7 

10t> ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Louisiana: Natchitoches, G. F. Atkinson, 5118, 5119; St. Mar- 

tinville, A. B. Langlois, be. 
Ohio: Cincinnati, C. G. Lloyd, 1677, 4495, 4502. 
Wisconsin: Madison, C. J. Humphrey, 2508 (in Mo. Bot. Gard. 

Herb., 42927). 
Kentucky: Mammoth Cave, C. G. Lloyd. 
Missouri: Kennett, H. von Schrenk (in Mo. Bot. Gard. Herb., 

42996) ; Neeleyville, F. C. Dewart (in Mo. Bot. Gard. Herb., 
5132, 5135). 

Mexico: W. Trelease (in Mo. Bot. Gard. Herb., 5123); Guer- 

navaca, E. W. D. Holway. 

Honduras: P. Wilson, 180, 671, comm. by N. Y. Bot. Gard. 

Herb. 

Cuba: C. Wright, 197, 271 (in Curtis Herb.), and 521, the type 

of S. Sprucei (in Kew Herb.) ; Baracoa, L. M. Underwood 
& F. S. Earle, 796, 1068, comm. by N. Y. Bot. Gard. Herb.; 
Ceballos, C. J. Humphrey, 2722 (in Mo. Bot. Gard. Herb., 

8638). 

Porto Rico: Sauerce, Mr. & Mrs. A. A. Heller, 81+3, 882, comm. 

■ N. Y. Bot. Gard. Herb.; Luquillo Mts., P. Wilson, 203 

(in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 
56600). 

Guadeloupe: in de Thiimen, Myc. Univ., 2001. 
St. Kitts: N. L. Britton & J. F. Cowell, 502, comm. by N. Y. 

Bot. Gard. Herb. 

Jamaica: A. E. Wight, comm. by W. G. Farlow; Castleton 

Gardens, W. A. & E. L. Murrill, 113, comm. by N. Y. Bot. 
Gard. Herb.; Cinchona, W. A. & E. L. Murrill, 530, comm. 
by N. Y. Bot. Gard. Herb.; Moneague, W. A. Murrill, 
1140, comm. by N. Y. Bot. Gard. Herb.; Troy and Tyre, 
W. A. Murrill & W. Harris, 996, 1037, comm. by N. Y. 
Bot. Gard. Herb. 

Trinidad: Carengo, M. A. Carriker, comm. by W. G. Farlow, II. 
Grenada: Grand Etang, R. Thaxter, comm. by W. G. Farlow, 3. 
Venezuela: Margarita, A. F. Blakeslee, comm. by W. G. 

Farlow. 




46. S. versicolor (Swartz) Fries, Epicr. 547. 1838; Berkeley 
Ann. & Mag. Nat. Hist. I. 10: 382. pi. 11. f. 13. 1842; Sacc 



1920] __ 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 10/ 



mar 



Syll. Fung. 6: 561. 1888; Massee, Linn. Soc. Bot. Jour. 27: 

172. 1890; Lloyd, Myc. Writ, 4- Letter 46:3. 1913. 

Plate 5, fig. 47. 

Helvetia versicolor Swartz, Prodr. 149. 1788.— Thelephor a 
versicolor Swartz, Fl. Ind. Oc. 3: 1934. 1806; Fries, Syst. 
Myc. 1: 438. lS2l.—Stereum radians Fr.es, R. Soc. Sci. Up- 
sal. Actis III. 1: 110. 1851; Sacc. Syll. Fung. 6: 573. 1888; 
Massee, Linn. Soc. Bot, Jour. 27: 188. pl.7.f.6. 1900. 

Illustrations: Berkeley, loc. cit.; Massee, loc. cit. 

Type: authentic specimen in Herb, of Brit. Mus. according 

to Berkeley. 

Fructification coriaceous-rigid, very thin, sometimes buff- 
yellow, clothed with silky, villous fascicles all lying in a radiat- 
ing direction, becoming glabrous and shining and minutely 
radially ridged or lineate, wood-brown to cinnamon-brown, the 

jin entire, not complicate; in structure 300-400 n thick, 
composed of densely, longitudinally arranged hyphae 3-3| m in 
diameter; hymenium even, glabrous, cream-color to avellaneous; 
no colored conducting organs, gloeocystidia, nor cystidia; spores 
hyaline, even, 4-5 X 2-2 | p. 

Fructifications 1-2! cm. broad, 1^-4 en . long, often laterally 

confluent. 

On dead wood. Florida, West Indies, Mexico, Dutch Guiana. 

September to February. Probably common in Jamaica. 

S. versicolor is a species intermediate between S. lobatum and 
S. rameale; its fructifications are smaller than those of S. lobatum, 
thinner, more completely glabrous at length, with margin not 
normally lobed, and usually retaining attachment by a narrow, 
resupinate side of the pileus as well as by the umbo, in which 
respect there is resemblance to the middle stage of development 
of S.fasciatum; the radial arrangement of the hairs and villous 
fascicles on the upper surface of the pileus is a highly distinctive 
character, as first pointed out by Berkeley. The coloration and 
hairy covering of fructifications of S. versicolor are somewhat 
similar to these characters in S. rameale, but the fructifications 
of the former are not lobed and folded together laterally and 
crisped nor as slender as those of S. rameale, as pointed out by 
Fries in his description of his S. radians. S. versicolor was 
formerly confused with S. fasciatum, especially in American 




ICO (VoL ' 7 

IDS ANNALS OF THE MISSOURI BOTANICAL GARDEN 

literature; it is doubtful whether S. versicolor occurs in the 
United States except very rarely in Florida. 

Specimens examined: 
Florida: Dade County, /. K. Small, 7089, 7122 (in N. Y. Bot. 

Gard. Herb, and Mo. Bot. Gard. Herb., 56650, 56651); 

Eustis, Lake County, L. M. Underwood, 1377 (in N. Y. 

Bot. Gard. Herb., Burt Herb., and Mo. Bot. Gard. Herb., 
42764). 

Cuba: C. Wright, 291 (in Curtis Herb.); Ceballos, C. J. Humph- 
rey, 2740 (in Mo. Bot. Gard. Herb., 15720) ; San Diego de los 
Baiios, Bro. Leon, 4861 (in N. Y. Bot. Gard. Herb, and Mo. 
Bot. Gard. Herb., 56647). 

Porto Rico: Maricao, N. L. Britton, J. F. Cowell & S. Brown, 

4420 (in N. Y. Bot. Gard. Herb., Burt Herb., and Mo. Bot. 
Gard. Herb., 56574); Rio Piedras, J. R. Johnston, 129, 282 
(in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 
56648, 56641) ; Sierra de Naguabo, J. A. Shafer, 3211, 3692, 

(in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 
56653-56655) . 

Jamaica: Farr (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. 

Herb., 56640); Cinchona, L. M. Underwood, 3239 (in N. 
Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 56595), 
N. L. Britton, 295, 296 (in N. Y. Bot. Gard. Herb, and Mo. 
Bot. Gard. Herb., 56642, 56643), F. S. Earle, 409, comm. by 
N. Y. Bot. Gard. Herb., W. A. & E. L. Murrill, 526, 539, 
comm. by N. Y. Bot. Gard. Herb, and 473 (in N. Y. Bot. 
Gard. Herb., Burt Herb., and Mo. Bot. Gard. Herb., 
56644) ; John Crow Peak, L. M. Underwood, 2433, comm. 
by N. Y. Bot. Gard. Herb.; Monkey Hill, W. A. Murrill, 
814, comm. by N. Y. Bot. Gard. Herb.; Rose Hill, F. S. 
Earle, 50, 282, 305, comm. by N. Y. Bot. Gard. Herb.; 
Sir John Peak, E. G. Britton, 1212 (in N. Y. Bot. Gard. 
Herb, and Mo. Bot. Gard. Herb., 56641); Troy and Tyre, 
W. A. Murrill & W. Harris, 853, 856, 1036, 1048, comm. 
by N. Y. Bot. Gard. Herb. 

Montserrat: Soufriere, J. A. Shafer, 919 (in N. Y. Bot. Gard. 

Herb, and Mo. Bot. Gard. Herb., 56645). 
Grenada: Annandale, W. E. Broadway (in N. Y. Bot. Gard. 

Herb, and Mo. Bot. Gard. Herb., 56656); Grand Etang, 

R. Thaxter, comm. by W. G. Farlow, 10. 



1920) 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 169 



Mexico: Trap, de la Conception, Liebman, type of Stereum 

radians (in Herb. Fries); Jalapa, W. A. & E. L. Murrill, 
343, comm. by N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard. 
Herb., 55477). 



47. S. rameale Schweinitz, Naturforsch. Ges. Leipzig Schrift. 
1 : 106. 1822 (under B. Sterea of Thelephora). Plate 5, fig. 48. 

Thelephora hirsuta Fries, Elenchus Fang. 1: 178. 1828, 
but not of Syst. Myc. i:439. 1821. — T. hirsuta p ramealis 
Schweinitz, Am. Phil. Soc. Trans. N. S. 4: 167. 1832 
Stereum complicatum Fries, Epicr. 548. 1838; Sacc. Syll. Fung. 
6:579. 1888; Massee, Linn. Soc. Bot. Jour. 27:178. 1890. 
— S. radians of Morgan, Cincinnati Soc. Nat. Hist. Jour. 10: 
194. 1888, but not S. radians Fries. — Telephora lobata Bertolo- 
nii, Accad. Sci. Bologna Mem. I. 7: 360. pi. 19. f. e-g. 1856; 
Underwood & Earle, Ala. Agr. Exp. Sta. Bui. 80: 232. 1897 
Stereum Bertolonii Saccardo, Sacc. Syll. Fung. 11: 120. 1895. 

Illustrations: Berkeley & Broome, Linn. Soc. Bot. Trans. 
2: pi. 14. f. 12-14. 1883; Bertolonii, he. cit. 

Type: in Herb. Schweinitz and in Herb. Fries. 

Fructifications coriaceous, thin, rigid, effuso-reflexed, rarely 
resupinate, with the reflexed portion consisting of small, umbo- 
nate pilei, which are sometimes subdivided into 
lobes, the pilei or lobes drying folded together or 



sped, fibrose-strigose, becoming glabrous on the 



*^ 



m 



portion, shining, with innate fibers 




radiating from the base, cinnamon-buff to hazel, 
more or less zoned; hymenium even, glabrous, ^ Flg * 24 ; 
light buff to cream-buff; in structure 300-450 m Spores x 650. 
thick, composed of densely, longitudinally ar- 
ranged, hyaline hyphae 3-3^ n in diameter, colored conducting 
organs 3-3^ m in diameter occasionally present; no cystidia 
nor gloeocystidia; spores white in spore collection, even, 
slightly curved, 6X2-2£ M . 

Fructifications sometimes covering areas only 5-10 mm. in 
diameter, and gregarious, at other times irregularly confluent 
over areas up to 3 cm. broad and 10 cm. and more long; indi- 
vidual pilei 2-10 mm. broad, 3-10 mm. long. 

On dead twigs and stumps of oak and other frondose species. 



[Vol. 7 



170 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Canada, throughout the United States, except in the Rocky 
Mountain region, in Mexico and the West Indies. July to 
January. Common in the United States. 

S. rameale varies somewhat under the different conditions as 
to climate and substratum in the great extent of North America 
where it is our commonest species of Stereum. In the United 
States and Canada one will hardly go amiss in referring to S. 
rameale any Stereum with numerous small pilei densely crowded 
together imbricately or laterally, strigose hairy near the region 
of attachment, and with marginal side shining, somewhat zonate, 
and pinkish buff to hazel in color, and with these pilei drying 
folded together along the sides, or radially plicate in a laterally 
confluent form. The pileus of S. rameale is thinner than that 
of S. hirsutum, only partially covered with hairs, which do not 
form as heavy a covering where present, and the pilei are folded 
together laterally and are smaller than those of S. hirsutum. S. 
sericeum has small, shining, very thin pilei between whitish and 
pale drab-gray on both surfaces — wholly lacking ruddy ochra- 
ceous coloration — and almost always growing on Carpinus 

caroliniana. 

Schweinitz communicated to Fries specimens of S. rameale 
which are still preserved in the herbarium at Upsala; Fries 
published the species as a synonym of S. hirsutum in Elenchus 
Fung.; Schweinitz yielded to the authority of Fries but pro- 
tested that S. rameale was a distinct variety, at least. Other 
American specimens of this species were received by Fries, who 
described and published them in 1838 as S. complicatum, over- 
looking the earlier and nearly identical specimens from Schwei- 
nitz and the earlier, appropriate name for the species. 

Specimens examined : 
Exsiccati: Bartholomew, Fungi Col., 2881, 4289, 4689, 4985; 

Ellis, N. Am. Fungi, 324; Ell. & Ev., Fungi Col., 307; 

Ravenel, Fungi Car. 2:30; Fungi Am., 117; Smith, Cent. 

Am. Fungi, 96, 97 — the latter under the name S. sericeum; 

de Thiimen, Myc. Univ., 1404. 
Canada, Ontario: Belleville, J. Macoun, 240; Port Credit, J. H. 

Fault, Univ. Toronto Herb., 317 (in Mo. Bot. Gard. Herb., 

44878); Toronto, R. P. Wodehouse, Univ. Toronto Herb., 

316 (in Mo. Bot. Gard. Herb., 44879). 






^ 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 171 



Maine: Oldtown, P. L. Richer. 

Vermont: Brattleboro, Grand View Mt., Lake Dunmore, Mid- 

dlebury, and Ripton, E. A. Burt. 

Massachusetts: Arlington, E. A. Burt; Amherst, P. J. Ander- 
son, 6 (in Mo. Bot. Gard. Herb., 55850); Cambridge, W. 
Trelease, 81 (in Mo. Bot. Gard. Herb., 5062); Stony Brook, 
E. A. Burt; Waltham, A. B. Seymour, 12 (in Mo. Bot. 
Gard. Herb., 22096); Wellesley, L. W. Riddle, 12; Wor- 
cester, G. E. Francis. 

Connecticut: C. C. Hanmer, 2075 (in Mo. Bot. Gard. Herb., 

43849); Mansfield, P. W. Graff, 12 (in Mo. Bot. Gard. 
Herb., 9854); New Canaan, P. Wilson, 63 (in Mo. Bot. 
Gard. Herb., 54739); South Windsor, C. C. Hanmer. 

New York: Sartwell (in Mo. Bot. Gard. Herb., 5062, 44235); 

Albany, H. D. House (in N. Y. State Mus. Herb, and Mo. 
Bot, Gard. Herb., 15954); Alcove, C. L. Shear, 1137, 1320, 
1323, 1331; Catskill Mts., C. H. Peck, in Ellis, N. Am. 
Fungi, 324; East Galway, E. A. Burt, three collections; 
Glasco, P. Wilson, 34, 37, 41, 57 (in Mo. Bot. Gard. Herb., 
54728, 54741, 54742, 54727); Ithaca, G. F. Atkinson, 190 
O. S., 2121, 7989, 22969, 22973-22975, C. J. Humphrey, 227, 
H. S. Jackson, Cornell Univ. Herb., 11375, 14376, W. A. 
Murrill, Cornell Univ. Herb., 3058, Van Hook, Cornell 
Univ. Herb., 7991, K. M. Wiegand, Cornell Univ. Herb., 
3258, L. A. Zimm, 83 (in Mo. Bot. Gard. Herb., 9064); 
Palisades, P. Wilson, 16, 21 (in Mo. Bot. Gard. Herb., 
54732, 54731); Scarsdale, Livingston d: Crane, comm. by 
N. Y. Bot. Gard. Herb., P. Wilson, 1, 26 (in Mo. Bot. Gard. 
Herb., 54737, 54730); West Fort Ann, S. H. Burnham, 15 
(in Mo. Bot. Gard. Herb., 44011); Williams Bridge, P. 
Wilson, 3, 31 (in Mo. Bot. Gard. Herb., 54740, 54729); 
Yonkers, P. Wilson, 1 (in Mo. Bot. Gard. Herb., 54727). 

New Jersey: Laning (in Mo. Bot. Gard. Herb., 5051, 44236, 

44238); Alpine, P. Wilson, 15, 9, 14, 5, 4 (in Mo. Bot. Gard. 
Herb., 54733-54736, 54738); Newfield, /. B. Ellis, in Ellis, 
Fungi Col., 307, and in de Thumen, Myc. Univ., 1404; 
New Brunswick, H. D. House (in N. Y. State Mus. Herb, 
and Mo. Bot. Gard. Herb., 54353). 

Pennsylvania: Bear Meadow, C. R. Orton & A. S. Rhoads. 13. 



•k 



■ 



(Vol. 7 



172 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



14 (in Mo. Bot. Gard. Herb., 44090, 44091); Bellefonte 
L. 0. Overholts, 3715 (in Mo. Bot. Gard. Herb., 54996) 



Kittannine, D. R. Sumstine, 8, 9, 12; North Garden, E 



Michener, 437 (in Mo. Bot. Gard. Herb., 44237) ; Shingleton 
Gap, A. S. Rhoads, 11 (in Mo. Bot. Gard. Herb., 44089); 
Spruce Creek, /. H. Faull, Univ. Toronto Herb., 313 (in 
Mo. Bot. Gard. Herb., 44885). 

Delaware: Newark, H. S. Jackson, B9. 

Maryland: Cabin John Bridge, C. L. Shear, 101*5; Cabin John 

Creek, A. S. Rhoads, coram, by L. 0. Overholts (in Mo. 
Bot. Gard. Herb., 55069); Chevy Chase, comm. by Mrs. 
F. W. Patterson (in Mo. Bot. Gard. Herb., 43730); Takoma 
Park, A. S. Rhoads, comm. by L. O. Overholts (in Mo. Bot. 
Gard. Herb., 55049), C. L. Shear, 1160. 

District of Columbia: Takoma Park, P. L. Richer, 818. 

Virginia: Mt. Vernon, P. L. Richer, 1121 in -part. 

North Carolina: Schweinitz, type (in Herb. Schweinitz and 

Herb. Fries); Chapel Hill, W. C. Coher, 8802, 2026, 1047, 
362, 383 (in Mo. Bot. Gard. Herb., 56657-56661); Salem, 
Schweinitz, the Thelephora ochroleuca of Schweinitz, Syn. 
N. Am. Fungi, 644 (in Herb. Schweinitz). 

South Carolina: H. W. Ravenel, in Ravenel, Fungi Car. 2:30; 

Clemson College, P. H. Rolfs, 1614, 1628; Davidson River, 
H. von Schrenh (in Mo. Bot. Gard. Herb., 42964); Society 
Hill, H. W. Ravenel (in Curtis Herb., 1439, under the name 

Stereum plicatum). 

Georgia: Atlanta, E. Bartholomew, 5674 (in Mo. Bot. Gard. 

Herb., 44217); Glenbrook Ravine, A. B. Seymour, from 
Farlow Herb., J (in Mo. Bot. Gard. Herb., 44649); Thom- 
son, H. H. Bartlett, comm. by W. G. Farlow. 

Florida: C. G. Lloyd, 4851, 4852; Camp Pinchot, W. H. Long, 

12212 (in Mo. Bot. Gard. Herb., 55143); Daytona, D. L. 
James, comm. by U. S. Dept. Agr. Herb.; Gainesville, 
H. W. Ravenel, in Ravenel, Fungi Am., 117; New Smyrna, 

C. G. Lloyd, 2112. 
Alabama: Dr. Gates, probably from the type collection of 

Telephora lobata Bertolonii, from Torrey Herb, (in N. Y. 
Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 56295); 
Auburn, F. S. Earle, four specimens in Burt Herb., and two 






1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 173 



others (in Mo. Bot. Gard. Herb., 5107, 56619 — the last in 
N. Y. Bot. Gard. Herb, also); Montgomery Co., R. P. 
Burke, 28 (in Mo. Bot. Gard. Herb., 17856). 

Mississippi: Biloxi, F. S. Earle, 29; Hattiesburg, C. J. Humph- 
rey, 5451; Jackson, E. Bartholomew, 5779, 5797, 5784 (in 
Mo. Bot. Gard. Herb., 44223-44225) and Bartholomew, 
Fungi Col., 4689; Laurel, C. J. Humphrey, 5430; Ocean 
Springs, F. S. Earle, 177 (in Mo. Bot. Gard. Herb., 5065). 

Louisiana: A. B. Langlois, 2906; Alden Bridge, W. Trelease 

(in Mo. Bot. Gard. Herb., 5047); Baton Rouge, C. J. 
Humphrey, 5699 (in Mo. Bot. Gard. Herb., 14102); New 
Orleans, E. Bartholomew, 5764 (in Mo. Bot. Gard. Herb., 
5440, 44222), E. A. Burt; St. Martin vrille, A. B. Langlois, 
be (in Burt Herb.), 1101 (in Mo. Bot. Gard. Herb., 5063); 
Shreveport, E. Bartholomew, in Bartholomew, Fungi Col., 
4689. 

Ohio: Cincinnati, A. P. Morgan, comm. by Lloyd Herb., 2633; 

College Hill, C. G. Lloyd, 1457; Linwood, C. G. Lloyd, 02833. 

Indiana: Avilla, W. H. Rankin (in Mo. Bot. Gard. Herb., 9183); 

Crawfordsville, D. Reddick, 12; Greer.castle, L. M. Under- 
wood (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 
56277). 

Illinois: Bowmansville, comm. by Univ. Wis. Herb., 4, and 

E. T. & S.A. Harper, 436; River Forest, E. T. & S. A. 
Harper, 709. 

Kentucky: Bowling Green, S. F. Price (in Mo. Bot. Gard. Herb., 

5036). 
Tennessee: Elkmont, C. H. Kauffman, 58, 61, 63 (in Mo. Bot. 

Gard. Herb., 16384, 3993, 1678) ; Nashville, E. Bartholomew, 

5634 (in Mo. Bot. Gard. Herb., 44214). 
Michigan: Chelsea, C. H. Kauffman, 23; New Richmond, C. H. 

Kauffman, 44, 43 (in Mo. Bot. Gard. Herb., 22507, 22856). 
Minnesota: E . L. Jensen, 2 (in Mo. Bot. Gard. Herb., 3939). 
Wisconsin : Miss A. D. Stucki, Univ. Wis. Herb., 7; Blue Mounds, 

Miss A. D. Stucki, Univ. Wis. Herb., 6; Madison, Miss A. 

D. Stucki, Univ. Wis. Herb., 10. 
Iowa: E. W. D. Holway. 
Missouri: B. M. Duggar, 568; Bismarck, L. O. Overholts (in 

Mo. Bot. Gard. Herb., 43701); Cox's Switch, H. von 






[Vol. 7 
174 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Schrenk (in Mo. Bot. Gard. Herb., 42892); Creve Coeur, 
E. A. Burt (in Mo. Bot, Gard. Herb., 44757); Columbia, 
L. E. Cline, comm. by B. M. Duggar, A555; Gasconade Co., 
W. Trelease (in Mo. Bot. Gard. Herb., 5128); Meramec, 
P. Spaulding (in Mo. Bot. Gard. Herb., 5019); Neeleyville, 
Dewart (in Mo. Bot. Gard. Herb., 5127, 5130); St. Francis 
River, W. Trelease (in Mo. Bot. Gard. Herb., 5129); St. 
Louis, E. A. Burt (in Mo. Bot. Gard. Herb., 8724, 44757), 
and H. von Schrenk (in Mo. Bot. Gard. Herb., 42873); 
Williamsville, B. M. Duggar & H. S. Reed, 47. 

Arkansas: Arkadelphia, L. M. Underwood (in N. Y. Bot. Gard. 

Herb, and Mo. Bot. Gard. Herb., 56620); Batesville, E. 
Bartholomew, in Bartholomew, Fungi Col., 2881; Cass, 
W. H. Long, 19835 (in Mo. Bot. Gard. Herb., 6384); 
Womble, W. H. Long, 19671, 19649, 19865 (in Mo. Bot. 
Gard. Herb., 6386, 6385, 8887); Wynne, W. Trelease (in 
Mo. Bot. Gard. Herb., 5039). 

Texas: H. W. Ravenel, 40 (in U. S. Dept. Agr. Herb.); Joaquin, 

E. Bartholomew, in Bartholomew, Fungi Col., 4985; Somer- 
ville, H. von Schrenk, 1. 

Colorado: Tolland, L. 0. Overholts, 2000 (in Mo. Bot, Gard. 

Herb., 54872). 

British Columbia: Hastings, J. Macoun; Sidney , J '. M acoun , 1 4 , 

382 (in Macoun Herb.) and 56, 72 (in Mo. Bot. Gard. Herb., 
5738, 5748). 

Washington: Bellingham, J. R. Weir, 543, 547, 593 (in Mo. 

Bot. Gard. Herb., 18629, 18712,36745); Metaline Falls, 
J. R. Weir, 5245, 590 (in Mo. Bot. Gard. Herb., 55650, 
36744); Seattle, W. A. Murrill, 137, comm. by N. Y. Bot. 
Gard. Herb, (in Mo. Bot. Gard. Herb., 55736). 

Oregon: Corvallis, W. A. Murrill, 892b, comm. by N. Y. Bot. 

Gard. Herb, (in Mo. Bot. Gard. Herb., 55719), and C. E. 
Owens, 2033, 2134, 2147 (in Mo. Bot. Gard. Herb., 43873, 
44697, 9186). 

California: R. A. Harper, 121, 128 (in N. Y. Bot, Gard. Herb. 

and Mo. Bot. Gard. Herb., 56621, 56622); Palo Alto, 
W. A. Murrill & L. S. Abrams, 1170, comm. by N. Y. Bot. 
Gard. Herb, (in Mo. Bot. Gard. Herb., 55710). 

Mexico: Jalapa, W. A. & E. L. Murrill, 57, 70, 348, comm. by 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 175 



N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 23108, 
3732, 54475), and C. L. Smith, in Smith, Central Am. Fungi, 
96, 97; Orizaba, W. A. & E. L. Murrill, 799, comm. by 
N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 54624) ; 
Trap, de la Conception, Liebman, authentic specimen of 
Stereum complicatum (in Herb. Fries). 

Porto Rico: Indiera Fria, N. L. Britton, J. F. Cowell & S. 

Brown, 4483 (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. 
Herb., 56623). 

Jamaica: Abbey Green, W. Harris, 1022; Cinchona, F. S. 

Earle, 360, and W. A. & E. L. Murrill, 600, both numbers 
comm. by N. Y. Bot. Gard. Herb.; Hope, F. S. Earle, 119, 
comm. by N. Y. Bot. Gard. Herb.; New Haven Gap, W. A. 
& E. L. Murrill, 770, comm. by N. Y. Bot. Gard. Herb.; 
Monkey Hill, W. A. Murrill, 790, 802, comm. by N. Y. Bot. 
Gard. Herb.; Rose Hill, F. S. Earle, 309, 312, comm. by 
N. Y. Bot. Gard. Herb. 



48. S. sericeum Schweinitz, Naturforsch. Ges. Leipzig Schrift. 
1 : 106. 1822 (in B. Sterea of Thelephora) ; Morgan, Cincinnati 
Soc. Nat. Hist. Jour. 10: 195. 1888; Sacc. Syll. Fung. 6: 579. 
1888. Plate 5, fig. 49. 

Thelephora striata Fries, Elenchus Fung. 1 : 178. 1828; 
Schweinitz, Am. Phil. Soc. Trans. N. S. 4: 167. 1832. — Ster- 
eum striatum Fries, Epicr. 548. 1838, but not of p. 551 nor of 
Hym. Eur. 641. 1874. 

Illustrations: Hard, Mushrooms, 456. text f. 383. 

Type : not found by me in Herb. Schweinitz although studied 
by Berkeley & Curtis, Acad. Nat. Sci. Phila. Jour. 3 : 220. 1856. 

Fructifications coriaceous, small, very thin and papery, ef- 
fuso-reflexed, laterally confluent, with reflexed portion divided 
into small pilei, sometimes orbicular and attached by a central 
point with margin free all around, the upper side 
whitish to cartridge-buff, shining, silky, with ^-y 

minute radiate fibrils, the margin entire, thinning ^ ^^^ 
to subfimbriate, not complicate; hymeniumeven, 
wood-brown when most deeply colored, becoming 
bleached ; in structure 250-300 p thick, composed « Flg : 25, 
ol densely and longitudinally arranged hyaline Spores x 665. 






[Vol. 7 



176 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



hyphae 3-3 



\ M 



in diameter; no colored conducting organs, 



gloeocystidia, nor cystidia present; spores hyaline, even, flat- 
tened on one side, 6-7|X3-3§ \i. 

ons \-\\ cm. in diameter, confluent along limbs 



Fructificat 



10 mm. broad, 3-10 mm 



10 cm. and more, the reflexed portion 5- 

long. 

In swampy woods on under side of dead twigs of Carpi 



orded rarely on Liquidambar and Ny 



ada to Louisiana 
Throughout the v 



d westward to M 
Infrequent. 



and 



;. Can 
Mexico 



Stereum sericeum is very appropriately named, for its 
to pale gray pilei are noteworthy by their silky or satiny 
they are smaller, thinner, and more flexible than those 
rameale and with innate rather than fibrose-strigose 



of S 
fibrils 



these pilei lack the ruddy and ochraceous hues characteristic 
of S. rameale; furthermore the pilei of S. sericeum are plane, 
while those of S. rameale are folded laterally or crisped. Never- 

of S. rameale 



theless I ha\ 



some 



specimens 



from the West and South which were sparsely developed and 



bleached out so as to simulate S 



In New England 



and New York, S. sericeum has been invariably 



Carp 



caroliniana when the substratum has been recorded, but else 

where S. rameale has sometimes been recorded on other substrata 

The concept of S. sericeum is that held by all American my 



cologists and is in conformity 



the specimens in Curtis 



Herbarium determined by Berkeley and Curtis who studied the 
authentic specimen. 

Specimens examined: 
Exsiccati: Ellis. N. Am. Fung., 19; Ell. & Ev., Fungi Col., 705; 



Ravenel, Fungi C 



Shear. N 



Ontario : London, J. Dearness : Ottawa, J 



H. Graham, Univ. Toronto Herb., 675 (in M 



Card. Herb.. 44918), and 



Lanaton. Univ 



Herb., 518, 594 (in Mo. Bot. Card. Herb 
Vermont: Middlebury, E. A. Burt, five coll 
Massachusetts: Wayland, A. B. Seymour, 



T23 (in M 



Bot. 



Herb 



C 



Gosh 



L. M. Underwood, 224 



N. Y. Bot 



Gard. Herb, and Mo. Bot. Gard. Herb 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 177 



New York: Sartwell (in Mo. Bot. Gard. Herb., 5045); Alcove, 

C. L. Shear, 1047, 1124, 1211, 1314, 1325, 1332, and in 
Shear, N. Y. Fungi, 312; Glasco, P. Wilson, 36 (in Mo. 
Bot. Gard. Herb., 54744); Grand View, H. von Schrenk 
(in Mo. Bot, Gard. Herb., 42795); Ithaca, G. F. Atkinson, 



178 0. S., 2827, 22968, and W. C. Mucnscher, 4 (in Mo. Bot. 

Gard. Herb., 56594); McLean, W. C. Muenscher, 98 (in 

Mo. Bot. Gard. Herb., 56596); Taughannock Gorge, W. C. 

Muenscher, 199 (in Mo. Bot. Gard. Herb., 56595). 
New Jersey: Newfield, J.B. Ellis, in Ellis, N. Am. Fungi, 19, E1L 

& Ev., Fungi Col., 705, and (in Mo. Bot. Gard. Herb., 5103). 
Pennsylvania: E. Michener, 399 (in Mo. Bot. Gard. Herb., 

5104); State College, L. O. Overholts, 3054 (in Mo. Bot. 

Gard. Herb., 5688). 
District of Columbia: Takoma Park, C. L. Shear, 957. 
North Carolina: Chapel Hill, W. C. Coker, 1043 (in Mo. Bot. 

Gard. Herb., 56668). 
South Carolina: Black Oak, H. W. Ravenel, in Ravenel, Fungi 

Car. i : 31. 
Florida: Tallahassee, comm. by W. G. Farlow. 
Alabama: Auburn, F. S. Earle (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56661-56663); Fayette Co., P. V. 

Diggers, comm. by A. H. W. Povah, 17 (in Mo. Bot. Gard. 

Herb., 20803); Montgomery Co., R. P. Burke, 32, 137 (in 

Mo. Bot. Gard. Herb., 15929, 10934); Tuskegee, C. W. 

Carver, 369 (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. 

Herb., 56664). 

Mississippi: Biloxi, F. S. Earle, 27. 

Louisiana: New Orleans, F. S. Earle (in N. Y. Bot. Gard. Herb. 

and Mo. Bot. Gard. Herb., 56660). 

Ohio: Cleveland, H. C. Beardslee; Columbus, W. A. Kellerman, 

in Kellerman, Ohio Fungi, 139 (in Mo. Bot. Gard. Herb., 
5042); Norwood, C. G. Lloyd, 2270; Oberlin, and also 
Penfield, F. D. Kelsey (in Mo. Bot. Gard. Herb., 56665 and 
56666 respectively). 

Indiana: Scottsburg, J. R. Weir, 5803 (in Mo. Bot. Gard. Herb., 

55643). 

Michigan: Agricultural College, Hicks, comm. by W. G. Farlow. 

Missouri: Columbia, B. M. Duggar, 553. 

7 



1 7Q ,V0L - 7 

178 ANNALS OF THE MISSOURI BOTANICAL GARDEN 






Mexico: Jalapa, W. A. & E. L. Murrill, 343 in part, coram 
by N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 56672) 



49. S. pubescens Burt, n. sp. Plate 5, fig. 50. 

Type: in Mo. Bot. Gard. Herb., N. Y. Bot. Gard. Herb., and 
Burt. Herb. 

Fructification coriaceous, thin, orbicular, conchate-reflexed, 
attached by one side and the center, reflexed all around but more 
broadly on the upper side, white, pubescent with soft matted 
hairs, not zonate nor sulcate; hymenium drying even or some- 
what radiately rugose, sorghum-brown to dusky drab, shining; 
in structure 600 ^ thick exclusive of the tomentum, with the 
occasional hy menial wrinkles standing out up to 120 /* further; 
intermediate layer bordered next to the tomentum by a narrow, 
dense, colored zone and composed of longitudinally arranged 
and somewhat loosely interwoven hyaline, thick-walled hyphae 
3| /x in diameter; no vesicular organs, conducting organs, 
gloeocystidia, nor cystidia present; hymenium composed of a 

single layer of simple basidia with 4 sterigmata; spores hyaline, 
even, oval, 6X4 n. 

Fructifications 3-10 mm. in diameter, reflexed 1-3 mm. 

On dead limbs of a frondose species. Montana. April. 
Probably rare. 

S. pubescens has small fructifications with some resemblance 
in aspect to those of Cenangium furfuraceum but white and 
pubescent with soft matted hairs. Specimens from this gather- 
ing were communicated by Ellis, No. 7014, to Cooke and were 
regarded by Cooke as a young Stereum, related to Stereum 
purpureum and, perhaps, young specimens of this species. S. 
pubescens differs sharply from S. purpureum in having no pyri- 
form, vesicular organs. The specimens are so mature that 
many basidia bearing sterigmata are present and occasionally 
spores. In the smaller specimens the hymenium is even but 
in those 1 cm. in diameter some broad, obtuse, radiating wrinkles 
are present, which may necessitate the transfer of this species 
from Stereum when better known from future collections. 

Specimens examined: 
Montana: Sheridan, Mrs. L. A. Fitch, in Ellis Collection, 7014, 

type (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 

56784). 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 179 



50. S. conicum Burt, n. sp. Plate 5, fig. 51. 

Type: in Farlow Herb, and in Mo. Bot. Gard. Herb. 

Pileus coriaceous, small, rather thick, conical, sessile, attached 
by the vertex, villose, with some specimens whitish to pale 
olive-buff and others between wood-brown and Sayal-brown; 
intermediate layer not bordered by a dark zone, nearly colorless, 
containing many thick-walled and somewhat incrusted hyphal 
ends 15-25X6 n but no colored conducting organs; hymenium 
even, drab, without cystidia; spores hyaline, even, 4-4|X2| /*. 

Pileus 2-4 mm. in diameter, 2-4 mm. high, about f-^mm. 

thick. 

Singly on small, dead, frondose twigs. Cuba. 

If carelessly glanced at, specimens of this species might be 
referred to S. ochraceo-flavum, but in S. conicum each of the eight 
fructifications which I have seen is truly conical, pendant, and 
attached by its vertex, while the pilei of S. ochraceo-flavum, S. 
ochroleucum, etc., are reflexed; the hymenium of S. conicum is 
glabrous, while that of S. ochraceo-flavum contains even-walled, 
non-incrusted cystidia 20-25X4-6 n, protruding 15 n. S. coni- 
cum is noteworthy by the very numerous thick-walled and some- 
what incrusted hyphal ends which are present in its intermediate 
layer. On the hymenial side these bodies curve towards the 
hymenium but do not reach its surface; on the opposite side 
they curve to the upper surface of the pileus and protrude as 
incrusted hairs forming a part of the villose covering of the 
pileus, a structural feature suggestive of Cyphella. The speci- 
mens of S. conicum were collected by Charles Wright during 
his last trip to Cuba in about 1860 but were not sent to Berkeley 
and Curtis for study. 

Specimens examined: 
Cuba: Fungi Cubensis Wrightiani, 842, C. Wright, type, comm. 

by W. G. Farlow (in Mo. Bot. Gard. Herb., 43906 and in 
Farlow Herb.). 



51. S. vibrans Berk. & Curtis, Linn. Soc. Bot. Jour. 10: 332. 

1868; Sacc. Syll. Fung. 6:577. 1888. Plate 5, fig. 52. 

An Stereum cupulatum Patouillard in Diss, Fl. Crypt. Antilles 

Fr. 233. 1904? 

TvDe: in Curtis Herb, and Kew Herb. 



i on ( VoL - 7 

180 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Fructifications coriaceous, orbicular, and attached by the 
center, or fan-shaped and laterally confluent, lobed, the upper 
surface velvety hirsute on the region of recent growth, becoming 
somewhat glabrous in the older region near place of attachment, 
narrowly concentrically sulcate, somewhat zonate, snuff-brown, 
becoming Saccardo's umber; hymenium even, Saccardo's umber 



drab, somewhat 



th 



the intermediate layer connected with the hairy covering by 
blackish dense crust; hyphae of intermediate layer snuff-brown, 
blackening by action of dilute potassium hydrate, longitudinally 
arranged, thick-walled, 3^-4 n in diameter; hy menial layer 
simple; no colored conducting organs, cystidia, nor aculeate 
paraphyses; spores hyaline, even, 4-5X2^-3 p. 
Pileus 2-5 cm. in diameter. 

On logs. Cuba and Jamaica. October and November. 
Rare. 

S. vibrans is related to S. crassum but seems distinct by having 
smaller spores and a thin, blackish, horn-like crust under the 
hairy covering; the other histological details are very similar 
however. S. vibrans may be distinguished from the other 
species of the West Indies by its tobacco color, pruinose hymen- 
ium, and lack of cystidia, gloeocystidia, conducting organs, 
and bottle-brush paraphyses. S. papyrinum is of similar colora- 
tion, but is more spongy, has incrusted cystidia, and does not 
have its intermediate layer bordered above by a crust. 

Specimens examined: 
Cuba: C. Wright, 530, type (in Curtis Herb.). 
Jamaica: Rose Hill, F. S. Earle, 299, 303, comm. by N. Y. Bot. 

Gard. Herb. 



52. S. crassum Fries, R. Soc. Sci. Upsal. Actis III. i: 111. 



1851 

1888. 



Sacc. Syll. Fung. 6 : 582. 



Herb 



Fructification coriaceous, resupinate, effused, sometimes re- 
flexed, villose, blackening, the margin obtuse, determinate, 
paler; hymenium even, dark chestnut-brown; in structure 1000 
n thick, with intermediate layer not bordered by a darker denser 
zone or crust, composed of longitudinally and rather loosely 



1920] 



BURT— THELEPHORACEAE OF NORTH AMERICA. XII 181 

arranged, dark-colored, thick-walled, stiff hyphae 3§-4i » in 
diameter, not incrusted, which give their color to the fructifica- 
tion; no colored conducting organs, gloeocystidia, nor cystidia; 
spores hyaline, 9X4 /x. 

According to the original collection of S. crassum in Herb. 
Fries, this is a very distinct species, characterized by very dark 
color throughout and by absence of colored conducting organs, 
cystidia, and gloeocystidia. It is probably of local distribution,' 
for I have seen but one collection which is even doubtfully 
referable to S. crassum. This specimen, collected at Motzo- 
rongo, is wholly resupinate, with hyphae dark-colored and 

obliquely from the substratum instead of running 



endi 



longitudinally, and the hymenium has dried pinkish buff 
Specimens examined: 

Mexico: Mirador, Liebman, type (in Herb. Fries); Motzorongo, 

near Cordoba, W. A. & E. L. Murrill, 985 (in N. Y. Bot! 
Gard. Herb, and Mo. Bot. Gard. Herb., 54648). 

53- S. radiatum Peck, Buffalo Soc. Nat. Hist. Bui. i : 62. 

1873; N. Y. State Mus. Rept. 26:72. 1874; Sacc. Syll. 

1888; Massee, Linn. Soc. Bot. Jour. 27: 195. 

Plate 5, fig. 53. 
b. radiatum var. reflexum Peck, N. Y. State Mus. Rept. 
49:45. 1896; Sacc. Syll. Fung. 14: 217. 1900.— An Thele- 
phora (Stereum) corrugata Leveille, Ann. Sci. Nat. Bot. Ill 



571 



1890. 



5:150. 1846? 



N. Y. State Mus. Herb 



margin 






around, sometimes reflexed on the upper side, the reflexed 
tion becoming black above, velutinous, 
crisped, and somewhat lobed; hymenium 
uneven, not polished, marked with thick ^^ $ { 
ridges radiating from the center, Sudan- 
brown, rarely black when turned upward «„ 9ft c ,. , 

, j 1 , ,. ,. , . ^ rig. Jb. £>. radiatum. 

and exposed to direct sunlight and weather; Spores x 665. 

in structure 1000 M thick, composed of 
densely and longitudinally arranged, colored hyphae 3|-4 M in 
diameter, whose color is dissolved by dilute potassium hydrate 
solution; no cystidia; spores from spore collections white, 
even, slightly curved, 9-10X3*-4 u. 



[Vol. 7 



182 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Fructifications 2 cm. in diameter up to 10X3 cm.; reflexed 
portion 2-8 mm. broad. 

Under side of hemlock, spruce, and pine boards and logs and 
charred wood. Canada to Pennsylvania and westward to 
Montana; received also from Russia where growing on rotten 

wood in greenhouse. 

S. radiatum is readily recognized by its bright, ferruginous 
hymenium with shallow broad ridges radiating from the center 
to the margin, and by the black upper side of the pileus when 
reflexed. The general aspect, coloration, and color changes 
with KHO solution are suggestive of some species of Hymen- 
ochaete but no setae are present. I endeavored to have com- 
parison made with the type of Thelephora corrugata in Museum 
of Paris Herbarium but Patouillard could not find the specimen 

there. 

Specimens examined: 
Exsiccati: Ellis, N. Am. Fungi, 407. 
Russia: on rotting wood in a greenhouse, Janczewsky (in N. Y. 

Bot. Gard. Herb, and Mo. Bot, Gard. Herb., 6173). 
Ontario: Harraby, E. T. & S. A. Harper, 686. 
Vermont: Howe (in N. Y. Bot. Gard. Herb, and Mo. Bot. 

Gard. Herb., 5962); Lake Willoughby, W. G. Farlow; Mid- 

dlebury, E. A. Burt, four collections. 
Massachusetts: Cambridge, W. G. Farlow; Sharon, A. P. D. 

Piguet, comm. by W. G. Farlow, (in Mo. Bot. Gard. Herb., 

55002). 
New York: Albany, C. H. Peck, in Ellis, N. Am. Fungi, 407; 

Alcove, C. L. Shear, 1801; Freeville, G. F. Atkinson, Cornell 
Univ. Herb., 18185; Ithaca, C. 0. Smith, H. H. Whetzel, 
L. M. Wiegand, Cornell Univ. Herb., 8029, 13809, and 3254 

respectively. 
Pennsylvania: State College, L. O. Overholts, 2658 (in Mo. 

Bot. Gard. Herb., 5917); Trexlertown, W. Herbst. 

Michigan: Seney, C. J. Humphrey, 1848 (in Mo. Bot. Gard. 

Herb., 17766). 
Montana: Darby, J: R. Weir, 363 (in Mo. Bot. Gard. Herb., 

16472). 

54. S. patelliforme Burt, n. sp. Plate 5, fig. 54. 

Type: In Burt Herb. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 183 



Fructification coriaceous-fleshy, resupinate, the margin be- 
coming free or narrowly reflexed, hoary with a few short hairs, 
drying cinnamon to bone-brown, the margin entire; hymenium 
even, waxy, cracking in drying, drying cinnamon to bone- 
brown ; in structure 500-800 m thick, composed of longitudinally 
and densely arranged, hyaline hyphae 3-3 § n in diameter, with 
the intermediate layer not bordered on the upper side by a 
denser, darker zone; hair-like cystidia hyaline, cylindric, flex- 
uous, 50-60x5-6 /*, emerging up to 40 m but rarely present; 
basidia simple, with 4 sterigmata, often protruded; spores 
hyaline, even, 9-10X3-4 m, somewhat curved. 

Fructifications 3X2 mm., up to 25X3 mm., the margin free 
all around and rolled up 1-2 mm. 

On fallen branches of Acer, Quercus, and ether frondose species. 
Washington, California, and New Mexico. August to April. 
Rare. 

S. patelliforme differs from our other Stereums by being of 
more fleshy consistency and with a waxy hymenium. In these 
characters it approaches Corticium, but it has the longitudinal 
arrangement of hyphae characteristic of Stereum and the margin 
becomes narrowly reflexed. These characters separate S. patelli- 
forme from our other Stereums with the exception of S. pubescens, 
which is snow-white on the upper side with a thick covering 
of fine soft hairs, is more broadly reflexed, and has a somewhat 
radiately rugose hymenium. 

Specimens examined: 
Washington: Bingen, W. N. Suksdorf, 713, type, 752, 753, 884, 

917. 
California: Campo Mts., C. D. Orcutt, 2005, comm. by U. S. 

Dept. Agr. Herb. 
New Mexico: Ute Park, Colfax Co., P. C. Standley, 14735, 

comm. by N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 

44951). 

55. S. ochraceo-flavum Schweinitz in Peck, N. Y. State Mus. 
Rept. 22: 86. 1869; Morgan, Cincinnati Soc. Nat. Hist. Jour. 
10: 195. 1888; Sacc. Syll. Fung. 6: 576. 1888; Massee, Linn. 
Soc. Bot. Jour. 27: 184. 1890. Plate 5, fig. 55. 

Thelephora ochraceo-flava Schweinitz, A.m. Phil. Soc. Trans. 

N. 8.4:167. 1832. 



184 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Type: in Herb. Schweinitz and Curtis Herb. 

Fructification coriaceous, thin, small, effuso-reflexed, some- 
times confluent along branches, often conical and attached by- 
one side and the umbo and some- 
times only by the umbo, the upper 
side villose-tomentose, somewhat fur- 
rowed, white, weathering gray; in 
structure 200-300 n thick below the 
hairy covering, with intermediate 
layer becoming bordered on the 

■p. o- o , „ upper side by a denser or colored 

Fig. 27. S. ochraceo-flavum. ■. i 

Hymenium showing three cys- zone when old an d weathered, com- 
posed of densely and longitudinally 
arranged, hyaline hyphae 3-4 /* in 




tidia, X 488. 



diameter; no colored 



conducting 



organs; hymenium even, 



"yellow," becoming cream-buff in the herbarium; cystidia not 
incrusted, obtuse, 20-25X4-6 n, protruding up to 15 n\ spores 
not found. 



Reflexed 



mm. broad, and about as lone: scattered 



conical pilei 3-5 mm. in diamet 
On dead branches of frondose 



Canada 



and 
July 

S, 



ard to Missouri, and in California and Mexico. 



■flavum may be 



ed at sight by 



white, conical fructifications heavily clothed with long, soft 



hairs and by its bright y 
cystidia afford a good d 



h} 



microsco 



The non-incrusted 
)ical character for 
3ecimens of S. sul- 
sted beyond their 
normal season of active growth, the upper side of the interme- 
diate layer becomes hardened and Dale golden. 



phuratum. In specimens which h 



Specimens examined : 
csiccati: Ellis, N. Am 



& Ev., Fund Col 



Ravenel, Fungi Am., 787; Ravenel, Fungi Car. 2: 31; 



de Thiimen, Myc. Uni\ 
Ontario: Ottawa, J. Macou 
Vermont: Middlebury, E. 1 
Massachusetts: D. W. Wei 

Mo. Bot. Card. Herb. 

Magnolia, W. G. Farloh 



comm. by C. G. Lloyd, 145 (in 
56687); Cambridge, E. A. Burt: 






1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 185 



Connecticut: Storrs, A. E. Moss, comm. by P. W. Graff, 38 

(in Mo. Bot. Gard. Herb., 44792). 
New York: Albany, H. D. House (in N. Y. State Mus. Herb., 

and Mo. Bot. Gard. Herb., 55209); East Galway, E. A. 

Burt; Ithaca, Cornell Univ. Herb., 219; Poughkeepsie, 

W. R. Gerard, 228, 261 (in N. Y. Bot. Gard. Herb.); Staten 

Island, L. M. Underwood (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56701). 
New Jersey: Newfield, J. B. Ellis, in Ellis, N. Am. Fungi, 17, 

Ell. & Ev., Fungi Col., 6, and de Thumen, Myc. Univ., 10. 
Pennsylvania : Bethlehem, Schweinitz, type (in Herb. Schweinitz 

and in Curtis Herb.); State College, J. F. Adams, 8 (in 

Mo. Bot. Gard. Herb., 44085). 
Maryland: Seven Locks, P. L. Richer, 1005; Takoma Park, 

C. L. Shear, 1119, 1240. 
Virginia: Park Lane, W. H. Long, 18463 (in Mo. Bot. Gard. 

Herb., 55101). 
North Carolina: Blowing Rock, G. F. Atkinson, 4816. 
South Carolina: H. W. Ravenel, in Ravenel, Fungi Car. 2: 31 

Summerville, C. L. Shear, 1228. 
Georgia: Darien, H. W. Ravenel, in Ravenel, Fungi Am., 787 

Fullerton, P. L. Richer, 918. 



Florida: C. G. Lloyd, 4859; Hanosassa (in N. Y. Bot. Gard 



Herb, and Mo. Bot. Gard. Herb., 56688); New Smyrna 



' C. G. Lloyd, 2089; Tampa, N. L. & E. G. Britton & J. A 
Shafer, 46 (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard 

Herb., 56689). 
Alabama: Auburn, F. S. Earle & C. F. Baker (in Burt Herb. 

and Mo. Bot. Gard. Herb., 5089) ; Montgomery Co., R. P. 

Burke, 22 (in Mo. Bot. Gard. Herb., 12291). 
Mississippi: Ocean Springs, F. S. Earle, 180 (in Mo. Bot. Gard. 

Herb., 5090). 
Michigan: New Richmond, C. H. Kaufman, 87 (in Mo. Bot. 

Gard. Herb., 44995). 
Wisconsin: Palmyra, Miss A. O. Stucki, Lniv. Wis. Herb., 40. 
Indiana: Millers, E. T. & S. A. Harper, 938. 
Tennessee: Elkmont, C. H. Kauffman, 59 (in Mo. Bot. Gard. 

Herb., 44971). 
Iowa: Decorah, E. W. D. Holway. 



[Vol. 7 
186 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Missouri: Allenton, Letterman, 48 (in Mo. Bot. Gard. Herb., 

5041). 

Arkansas: Cass, W. H. Long, 19833 (in Mo. Bot. Gard. Herb., 

17807). 

California: Campo Seco, W. H. Thomas, 3 (in N. Y. Bot. Gard. 

Herb, and Mo. Bot. Gard. Herb., 86690). 

Mexico: Jalapa, W. A. & E. L. Murrill, 347, comm. by N. Y. 

Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 54468); Ori- 
zaba, J. G. Smith, 511 (in Mo. Bot. Gard. Herb., 437). 



56. S. abietinum Persoon, Myc. Eur. 1 : 122. 1822 (under **** 
Stereum of Thelephora) ; Fries, Obs. Myc. 1: 274. 1815, and 
ed. 2, 1824; Epicr. 552. 1838; Hym. Eur. 643. 1874; Sacc. 



Syll. Fung. 6: 574. 1888. 



fig 



Thelephora abietina Persoon, Syn. Fung. 573. 1801; Fries, 
Syst. Myc. 1 : 442. 1821. — Hymenochaete abietina (Pers.) Mas- 
see, Linn. Soc. Bot. Jour. 27: 115. 1890. — Thelephora stri- 
ata Schrader, Spic. Fl. Germ. 186. 1794. — Stereum striatum 
Schrader ex Fries, Epicr. 551. 1838; Hym. Eur. 641. 1874; 
Sacc. Syll. Fung. 6:565. 1888. — Lloydella striata (Schrad.) 
Bresadola in Lloyd, Myc. Writ. 1. Myc. Notes 6:51. 1901. 

Stereum glaucescens Fries, Hym. Eur. 644. 1874; Sacc. Syll. 
Fung. 6: 575. 1888. — Hymenochaete fimbriata Ellis & Ever- 
hart, Jour. Myc. 1 : 149. 1885; Sacc. Syll. Fung. 6: 599. 1888; 
Massee, Linn. Soc. Bot. Jour. 27: 113. 1890. — Hymenochaete 
abnormis Peck, N. Y. State Mus. Rept. 42 : 126. pi. l.f. 13-16. 
1889; Sacc. Syll. Fung. 9: 227. 1891. 

Illustrations: Istvanffi, Jahrb. f. wiss. Bot. 29: pi. 5. f. 16, 17; 
Patouillard, Essai Tax. Hym. 72; Peck, N. Y. State Mus. Rept. 
42: pi l.f. 13-16. 

Fructification coriaceous-spongy, dry, thick, resupinate, 
effused, rarely reflexed, with upper side tomentose, obscurely 
zonate, burnt umber, tuberculate or uneven; hymenium varying 
from light drab to cinereous or glaucous; in structure 400-900 m 
thick, of which the intermediate layer and the hymenium 
together constitute 300-600 m; intermediate layer composed of 
longitudinally arranged and interwoven colored hyphae 3-3^ 
m in diameter and bordered on its outer side by a darker, denser 
zone which connects it with the tomentose covering; hymenial 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



187 



layer becoming zonateand con- 
taining numerous colored cys- 
tidia having more or less the 
appearance of colored conduct- 
ing organs; cystidia colored, 
cylindric, obtuse, even, rough- 
walled or more or less incrusted, 



90 
60 



M 



> 



6-8 n, protruding up t< 
res hyaline, even, flat 



tened on one side, 9-13 X 
Resupinate specimens 

2-5 cm., reflexed margi 

mm. broad. 

On wood and logs of 



n 



5 



8X 
3-8 









S 



Abies 



Fig. 28. 



S. 



abietinum.. Section X 



d P 



68; crust-like zone, z; hymenium con- 



Xew Hampshire taining colored cystidia, h; cystidium, c, 



to Washington and 
June to October. 



and spores, s, X 488. 



Rare. 



S. abietinum usually occurs resupinate, bu : its thick, separable, 
felty fructifications are suggestive of a resupinate Stereum, and 
this view is confirmed by the presence of the intermediate layer 
when radial, vertical sections are examined. The cinereous, 
pruinose surface of the hymenium due, however, to whitish, 
cobwebby filaments rather than powdery grains, is highly char- 
acteristic and shared only by the western S. rugisporum, as 
are also the colored cylindric cystidia. S. rugisporum is sepa- 
rated by its odor of anise, much thicker and more broadly 
reflexed pilei, and presence in occasional collections of colored 
spores imbedded in the deeper zones of the hymenium. 

I have included Hymenochaete fimbriata among the synonyms 
of S. abietinum, but it may prove to belong with S. rugisporum 

instead. 

Specimens examined: 
Exsiccati: de Thiimen, Myc. Univ., 1107. 
Norway: Christiania, M. N. Blytt, type of Stereum glaucescens 

(in Herb. Fries). 
Sweden: Stockholm, L. Romell, 29; Upsali, C. G. Lloyd, 08521 

(in Lloyd Herb, and Mo. Bot. Gard. Herb., 55497). 
Finland: Mustiala, P. A.Karsten, in de Thiimen, Myc. Univ., 

1107. 



188 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



Italy (?) : locality not stated, G. Bresadola. 

New Hampshire: Crawford Notch, L. 0. Overholts & A. S 

Rhoads (in Mo. Bot. Gard. Herb., 56342); North Conway 
L. 0. Overholts, 4558 (in Mo. Bot. Gard. Herb., 55633). 
nont: Smugglers Notch, Mt. Mansfield, E. A. Burt. 



Vei 



New York: Cascadeville, Adirondack Mts., C. H. Peck, type of 

Hymenochaete abnormis (in N. Y. State Mus. Herb.). 
Wisconsin: Madison, M. C. Jensen, comm. by C. J. Humphrey, 



618. 
Montana: Yell 



fimbriata from J. B. Ellis (in Kew Herb 
Canada: Rocky Mts., Lake O'Hara, /. M 
Washington : Mt. Paddo, W. N. Suksdorf. 



Park, part of type of Hymenochaete 



2. 



57- S. rugisporum (Ell. & Ev.) Burt, n. comb. 



Hymenochaete 



Phila. Proc. 1890: 219. 1890; Sacc. S;y 
Type: in N. Y. Bot. Gard. Herb 



Plate 6, fig. 58. 
& Everhart, Acad. Nat. Sci. 

11. Fung. o:228. 1891. 



ficati 



coriaceous-spongy, dry, thick 



reflexed 



finally umbonate along line of attachment to substratum, the 

upper side tomentose, concentrically sulcate, 
snuff-brown when young and remaining so 
on the obtuse margin, elsewhere weathering 
neutral gray, with an anise-like odor in the 
herbarium; hymenium even, light mouse- 




becoming light drab 



3 



mm. thick, with intermediate layer and hy 



menium together 8( 
intermediate layer 



thick and the 

d with the 



Fig. 29. S. 



mentose 



porum 



r ugis- 
Portion of see- 



by a dark 



zone, 



hyphae of the 



tion x 488, showing col- m *ermediate layer colored, 2-4 M in diame 



ored imbedded spores. 



hymenial layer becomin 



itudinally arranged 
woven, curving outward into 



hymenial 



olored cystidia and sometimes 



to 1000 n thick, zonate, 



X3-3| fx, even 



■9 



h-walled; cystidia colored, cylindric 



1 



ugh or granule-incrusted 



150X7-9 ft, pro- 



1920] 



same 



BURT— THELEPHORACEAE OF NORTH AMERICA. XII 189 

truding up to 120 m, starting from all parts of the hymenial 
layer and subhymenium; basidiospores as seen on basidia, 
hyaline, even, 9-13X3-4 a M , borne 4 to a basidium. 

Reflexed portions 1-4 cm. long and wide, sometimes laterally 
confluent for 6-8 cm.; resupinate parts 3f about the 
dimensions. 

On dead Abies, Picea, Pinus, and Larix. In Rocky Mt. 
states and British Columbia to Arizona. July to September. 

Reflexed specimens of S. rugisporum may be recognized by 
their thick, felty, or spongy pilei, deeply concentrically sulcate, 
and snuff-brown or partly gray in color, with a whitish, pruinose 
hymenium, and an odor of anise; collections so far made indicate 
that this species is restricted to conifers of mountainous regions. 
Microscopic examination of sections shows characteristic cy- 
lindric, colored cystidia, which in the subhymenium and the 
deeper zones of the hymenium are not readily distinguishable 
from such colored conducting organs as occur in many species 
of Stereum. There is, however, no record of bleeding from 
wounds of the hymenium of S. rugisporum and S. abietinum. 
The type specimen of S. rugisporum contains colored spores, 
usually even, but occasionally rough-walled, imbedded in the 
deeper zones of the hymenium; similar spores occur in some, but 
not all, of the collections cited below, but the collections are so 
similar in other characters that I regard these colored imbedded 
spores as an important, occasional character of the species, which 
will positively identify some collections. 

The type of Hymenochaete fimbriate, was collected in Yellow- 
stone Park, Montana, on Pinus Murray ana; the specimen is 
wholly resupinate and does not show colored, imbedded spores 
in the preparations which I preserved. I regarded this specimen 
as not specifically distinct from S. abietinum, but the type 
station of H. fimbriata makes me uncertain as to whether the 
latter may not yet be demonstrated to be resupinate S. rugis- 
porum instead. When so demonstrated, the specific name 
fimbriatum should be used for the species because of earlier 
publication. 

Specimens examined: 

Wyoming: Fox Park, /. R. Weir, 10009 (in Mo. Bot. Gard. 

Herb., 55788). 



[Vol. 7 



190 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Colorado: Silverton, E. R. Hodson, comm. by C. J. Humphrey, 

1551; Tolland, L. 0. Over holts, 1781, 2336 (in Mo. Bot. 

Gard. Herb., 56042, 56761); Yankee Doodle Lake, F. J. 

Seaver & E. Bethel (in N. Y. Bot. Gard. Herb, and Mo. Bot. 

Gard. Herb., 56729). 
Idaho: Bonanza, G. G. Hedgcock, comm. by C. J. Humphrey, 

2168 (in Mo. Bot. Gard. Herb., 10377); Coolin, J. R. Weir, 

11476 (in Mo. Bot. Gard. Herb., 56724); Leesburg, F. S. 

Wolpert, comm. by J. R. Weir, 7033 (in Mo. Bot. Gard. 

Herb., 55463); Priest River, E. E. Hubert, comm. by J. R. 

Weir, 11655 (in Mo. Bot. Gard. Herb., 56725). 
British Columbia: J. Macoun, 94, type (in N. Y. Bot. Gard. 

Herb.). 
Washington: Olympic Mts., T. C. Frye, 1 (in N. Y. Bot. Gard. 

Herb, and Mo. Bot. Gard. Herb., 56730); Seattle, W. A. 

Murrill, 130, 146 (in N. Y. Bot. Gard. Herb, and Mo. Bot. 

Gard. Herb., 56731, 56732) and /. M. Grant, 2066, comm. 

by C. G. Lloyd (in N. Y. Bot. Gard. Herb., 56728). 
Arizona: Agassiz, W. H. Long, 19445 (in Mo. Bot. Gard. Herb., 

44734); Mt. Humphrey, Flagstaff, W. H. Long, 21306- 

21308, 21310 (in Mo. Bot. Gard. Herb., 54897-54899, 
54901); Interior Basin, San Francisco Peaks, W. H. Long, 

21309. 21311 (in Mo. Bot. Gard. Herb., 54900, 54902). 



58. S. ambiguum Peck, N. Y. State Mus. Rept. 47: 14 
94; Sacc. Syll. Fung. 11: 122. 1895. Plate 5, fig. S 

Type: in N. Y. State Mus. Herb. 

Fructifications coriaceous, dry, resupinate, effused, rarely m 
wlv reflexed, with the upper side tomentose, Prout's brow 



margin 



brighter colored, antique brow 



determinate; hymenium velvety, raw umber to Saccardo's 
umber when mature and thick, becoming deeply cracked in 
drying; in structure 600-1400 n thick, with an intermediate 
layer 400-600 m broad, composed of longitudinally interwoven, 
colored hyphae 3-4 /x in diameter, and with a zonate hymenial 
layer up to 800 p thick containing colored incrusted cystidia in 
all the zones; sections darkened by KHO solution; cystidia 
colored, cylindric, obtuse, usually incrusted, 100-150X7-12 n, 
protruding up to 100 n', basidiospores white in spore collection, 



1920 J 



BURT— THELEPHORACEAE OF NORTH AMERICA. XII 



191 




&;■• 




^ 



s 



even, 10-13 X3§-4J M ; colored 
spores 12X3|-4 y. sometimes 
occur in deeper zones of the 
hymenium. 

Resupinate part 1-8 cm. long, 
1-4 cm. wide, reflexed part 1-5 
mm. broad in the only reflexed 
specimen known. 

On logs of Abies and, perhaps, 
Pinus Strobus. Vermont and 
New York. June to November. 
Very rare. 

8, ambiguum belongs in the 
group of species with S. abietinum 
and S. rugisporum on account 
of similarity in microscopic structure including the colored 
cystidia. It may be separated from both these species at sight 
by the color of its hymenium which is permanently umber and 
not at all cinereous nor glaucous. There is a difference in 




O 



Fig. 30. S. ambiguum. Section of 
hymenial re ;ion X 68 ; peripheral part 
of cystidiuni, c, and spores, s, X 650. 



chemical composition 
blackens the sections 



o, for dilute potassic hydrate 
d becomes itself discolored £ 



the 



case of species of Hymenochaete. In fact, the general aspect 



like that 



;, thick, mature, deeply cracked specimens 
Hymenochaete spreta — a species 



which 



exceptionally on a coniferous substratum 



possibl 



S. ambiguum occurs in reflexed form in the state of Washing- 
ton, for the collection cited under S. rugisporum, Olympic 



Mt 



C. Frye, 1, resembles S ... m 

;h condition for confident reference here 



Specimens examined: 

Vermont: Middlebury, C. G. Lloyd, 10652 (in Lloyd Herb, and 

Mo. Bot. Gard. Herb., 44585); Ripton, E. A. Burt; Smug- 
glers Notch, Mt. Mansfield, E. A. Burt. 

New York: Adirondack Mts., C. H. Peck, type (in N. Y. State 

Mus. Herb.) ; Averyville, C. H. Peck (in N. Y. State Mus. 
Herb, and Mo. Bot. Gard. Herb.. 55699). 



50. S. umbrinum Berk. & Curtis, Grevillea 



1873 



Wakefield, Kew Bui. 101 «5 : 369 



1 9 15. — Com pare Stereum umbri 



[Vol. 7 



192 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



num Fries in Lehmann, Plantae Preissianae 2: 137. 1847. 

Plate 6, fig. 59. 

Thelephora crassa Leveille in Gaudichaud, Voyage Bonite Bot. 
i:190. pi. 139. f. 1. 1846. Not Stereum crassum Fries, R. 
Soc. Sci. Upsal. Actis III. 1: 111. 1851 .— H y menochaete 
crassa (Lev.) Berkeley in Cooke, Grevillea 8: 148. 1880; Sacc. 
Syll. Fung. 6: 597. 1888; Massee, Linn. Soc. Bot. Jour. 
27: 114. 1890.— H. umbrina Berk. & Curtis in Cooke, Gre- 
villea 8: 148. 1880; Morgan, Cincinnati Soc. Nat. Hist. 
Jour. 10: 198. 1888; Sacc. Syll. Fung. 6: 598. 1888; Massee, 
Linn. Soc. Bot. Jour. 27: 113. 1890.— //. vinosa (Berk.) 
Cooke, Grevillea 8: 149. 1880; Sacc. Syll. Fung. 6: 600. 
1888.— //. multispinulosa Peck, Bot. Gaz. 7: 54. 1882; Sacc. 
Syll. Fung. 6: 600. 1888; Massee, Linn. Soc. Bot. Jour. 27: 
108. 1890.— //. scabriseta Cooke in Ravenel, Fungi Am., 
717. 1882; Massee, Linn. Soc. Bot, Jour. 27: 113. pi. 5.J. 7. 
1890.— Lloydella scabriseta (Cooke) v. Hohn. & Litsch. K. Akad. 
Wiss. Wien Sitzungsber. 115: 1580. 1906.— Hy menochaete pur- 
purea Cooke & Morgan in Cooke, Grevillea 11: 106. 1883; 
Morgan, Cincinnati Soc. Nat. Hist, Jour. 10: 198. 18S8; Sacc. 
Syll. Fung. 6: 597. 1888; Massee, Linn. Soc. Bot. Jour. 27: 
115. 1S9Q .— Knieffia purpurea (Cooke & Morg.) Bresadola, 
Ann. Myc. 1 : 100. 1903. — Peniophora intermedia Massee, Linn. 
Soc. Bot. Jour. 25: 143. 1889; Sacc. Syll. Fung. 9: 238. 1891. 



H y menochaete Kalchbrenneri Massee, Linn. Soc. Bot, Jour. 27 

116. 1890; Sacc. Syll. Fung. 9: 230. 1891. 

Illustrations: Gaudichaud, Voyage Bonite Bot. pi. 139. f. 1; 
Linn. Soc. Bot. Jour. 27: pi. 5. f. 7. 

Type: in Kew Herb, and Curtis Herb. 

Fructifications coriaceous-spongy, resupinate, effused, often 
becoming reflexed, light vinaceous lilac to dark lavender when 
young, at length brownish drab to snuff-brown, the upper surface 
spongy, pitted, somewhat sulcate, the reflexed margin thick, 
entire; hymenium even, somewhat velvety, sometimes cracking 
in drying, light vinaceous lilac to snuff-brown; in structure 
500-1000 n thick, composed of loosely interwoven, slightly 
colored hyphae 3|-5 n in diameter, not forming an intermediate 
layer; in the subhymenial region thick-walled organs 5-6 n 
in diameter, darker colored than the hyphae, originate among the 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



193 



hyphae and curve outward 

hymenium 



h 



the 



as sharp-pointed cystidia, 

rough-walled, or in- 



) 



even, rouj 

crusted, 100-200X6-10 M 
protrudingupto^/z; spores 
white in spore collection, 

even, 6X3^ /*. 

Resupinate on are* 

cm. in diameter, bec< 

laterallv confluent for 



3 



15 



cm 




Fig. 31. S. vmbrinum. Section of hyme- 



, reflexed portion 2-5 
mm. broad. 

On fallen limbs of oak, 
hickory, and other fron- 

dose species. North Caro- n ; a i region X 488, showing z, cystidia. 

lina to Texas and south- 
ward from Ohio and Illinois, in Arizona, West Indies, and 
Central America; occurs also in Poland, Cochin China, and 
Australia. September to February, but collected occasionally 
in the other months of the year. 

S. umbrinum may be recognized by the purple color of young 
specimens which fades or changes finally to snuff-brown, al- 
though usually showing a vinaceous tinge, and by its remarkable 
cystidia, which, on account of their color £,nd lack of conspicuous 
incrustation, verge towards setae. However, these organs are 
paler colored and much more elongated *:han undoubted setae; 

furthermore, sections of fructifications in w T hich these colored 
cystidia are present do not immediately darken when dilute 
potassium hydrate is brought in contact with them, as inva- 
riably happens to sections containing true setae. It has seemed 
best to retain for this species the name Stereum umbrinum B. & 
C, because the type of Stereum umbrinum Fr., Herb. Preiss., 
No. 2686, collected in Australia on Banksia Menziesii, must be 
found and studied to complete the Friesian description before it 
can be known w r hether the Preiss specimen is not really a 
Hymenochaete, Eichleriella, Auricularia, or, perhaps, even identi- 
cal with S. umbrinum B. & C, a common species in Australia. 
The presence of a white, intermediate layer seems to preclude the 



s 



1ft i [Vol. 7 

iy4 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



latter possibility. No. 2686 has not been found in the Preiss 

series of specimens in the Missouri Botanical Garden Herbarium; 

perhaps it is most likely to be found in the Stockholm collection. 
Specimens examined: 

Exsiccati: Ellis, N. Am. Fungi, 606 b, under the name Stereum 

papyrinum, and 1108; Ell. & Ev., N. Am. Fungi, 2315; 
Ravenel, Fungi Car. 2 : 36, under the name S. papyrinum; 
Ravenel, Fungi Am., 118, under the name S. papyrinum, 
the type distribution of Peniophora intermedia, and 445, and 
717, the type distribution of Hymenochaete scabriseta; 
Rabenhorst, Fungi Eur., 3524; de Thumen, Myc. Univ., 
1504, under the name Corticium murinum, the type distribu- 
tion of Hymenochaete Kalchbr enneri . 

North Carolina: Asheville, E. Bartholomew, 5653 (in Mo. Bot. 

Gard. Herb., 44215); Creedmoor, J. G. Hall, comm. by 
Lloyd Herb., 10299 (in Mo. Bot. Gard. Herb., 55465). 

South Carolina: H. W. Ravenel, Curtis Herb., 1903, type (in 

Kew Herb.), and in Ravenel, Fungi Car. 2:36; Aiken, 

H. W. Ravenel, in Ravenel, Fungi Am., 445, and H. W. 

Ravenel, 1716 (in Curtis Herb., 2308, under the name 

Hymenochaete cervina); Clemson College, P. H. Rolfs, 1615, 
1633. 

Georgia: Darien, H. W. Ravenel, in Ravenel, Fungi Am., 117; 

Tallulah Falls, A. B. Seymour, comm. by W. G. Farlow, GG. 

Florida: C. G. Lloyd, 2134, 4857, and W. W. Calkins, in Ellis, 

N. Am. Fungi, 606 b; Eustis, R. Thaxter, 12 (in Farlow 
Herb, and Mo. Bot. Gard. Herb., 43931); Gainesville, N. 
L. T. Nelson, comm. by Lloyd Herb., 427 (in Mo. Bot. Gard. 
Herb., 55624), and H. W. Ravenel, in Ravenel, Fungi Am., 
118; Green Cove Springs, G. Martin, in Ellis, N. Am. Fungi, 

1108; New Smyrna, C. G. Lloyd, 192, 2122, 2134. 
Alabama: Peters, 770 (in Curtis Herb., under the name S. 

papyrinum); Auburn, P. H. Mell (in U. S. Dept. Agr. 
Herb, and Mo. Bot. Gard. Herb., 5106) ; Mobile, E. Barthol- 
omew, 5751 (in Mo. Bot. Gard. Herb., 44221); Montgom- 
ery, R. P. Burke, 139, 150 (in Mo. Bot. Gard. Herb., 
21228, 44906); Talapoosa region, F. S. Earle & C. F. Baker 
(in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb 
56598). 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 195 



Louisiana: A. B. Langlois, comm. by W. G. Farlow (in Mo. 

Bot. Gard. Herb., 44650); St. Martinville, A. B. Langlois, 
A, B, C, ag, and an unnumbered specimen and in Ell. & Ev., 
N. Am. Fungi, 2315. 

Ohio: A. P. Morgan, 11, type of Hymenochaete purpurea (in 

Kew Herb.); Cincinnati, C. G. Lloyd, 190, and A. P. 

Morgan, comm. by Lloyd Herb., 2626; Linwood, C. G. 
Lloyd, 2261. 

Indiana: Greenwood, M. C. Jensen, comm. bv C. J. Humphrey, 

2133 (in Mo. Bot. Gard. Herb., 22825). * 
Illinois: Christopher, C. J. Humphrey, 2133 (in Mo. Bot. Gard. 

Herb., 42926); Genesee, E. T. & S. A. Harper, 824. 
Missouri: Bismarck, L. 0. Overholts (in Mo. Bot. Gard. Herb., 

56716); Columbia, B. M. Duggar, 57 1; Pacific, L. 0. 

Overholts, 3162 (in Mo. Bot. Gard. Herb., 5718); Perryville, 

C. H. Demetrio, in Rabenhorst, Fungi Eur., 3524; Pickering, 

E. Bartholomew, 6424 (in Mo. Bot. Gard. Herb., 55194); 

St. Louis, N. M. Glatfelter, 1187, comm. by N. Y. Bot. Gard. 

Herb.; Valley Park, E. A. Burt (in Mo. Bot. Gard. Herb., 

44056, 44061). 

Arkansas: Bigflat, W. H. Long, 19858, 19895 (in Mo. Bot. Gard. 

Herb., 8965, 8883); Cass, W. H. Long, 19832, 19905 (in 
Mo. Bot. Gard. Herb., 8884, 8885); Womble, W. H. Long, 
19821 in part, 19869 (in Mo. Bot. Gard. Herb., 14650, 9142). 

Texas: Gillespie Co., C. Jermy, 444 (in Mo. Bot, Gard. Herb., 

5171); Gonzales, C. L. Shear, 1229. 

Arizona: 34 near Camp Lowell, C. G. Pringle, type of Hymeno- 
chaete multispinulosa (in N. Y. State Mus. Herb, and a 
portion in Burt Herb.). 

Cuba: C. Wright, Fungi Cubenses Wrightiari, 832, comm. by 

W. G. Farlow (in Mo. Bot. Gard. Herb., 43908), and C. G. 
Lloyd, 165 (in Mo. Bot. Gard. Herb., 55153); Ciego de 
Avila, Earle & Murrill, 607, comm. by N. Y. Bot. Gard. 
Herb.; La Magdalena, Earle & Baker, 2470, comm. by 
N. Y. Bot. Gard. Herb.; San Diego de Los Baiios, Earle 
& Murrill, 263, comm. by N. Y. Bot. Gaid. Herb. 

Porto Rico: Rio Piedras, J. A. Stevenson, 2389 (in Mo. Bot. 

Gard. Herb., 9441). 

Guatemala: Secanquim, W. R. Maxon & R. Hay, 3140a 



[Vol. 7 



196 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Cochin China: authentic specimen of Thelephora crassa from 

LeVeille (in Kew Herb.). 
Australia: W. N. Cheesman, comm. by E. M. Wakefield, Kew 

Herb, (in Mo. Bot. Gard. Herb., 44582); Victoria, J. G. 
Luehmann, in de Thiimen, Myc. Univ., 1504, under the 
name of Corticium murinum, the type distribution of Hy- 
menochaete Kalchbrenneri. 



6o. S. papyrinum Montagne in Ramon de la Sagra, Hist. 
Cuba PI. Cell. 374. 1842; ibid., folio ed., 9: 228. 1845; Syll. 
Crypt. 178. 1856; Berk. & Curtis, Linn. Soc. Bot. Jour. 10: 

Plate 6, fig. 60. 



331. 



1868. 




Peniophora papyrina (Mont.) Cooke, Grevillea8: 20. pi. 12^. 

9. 1879; Sacc. Syll. Fung. 6: 641. 1888; Massee, Linn. 
Soc. Bot. Jour. 25: 140. 1889. — Stereum nicaraguense Berk. 
& Curtis, Am. Acad. Arts & Sci. Proc. 4: 123. 1853; Sacc. 
Syll. Fung. 6: 567. 1888. — S. nicaraguae Berk. & Curtis in 
Massee, Linn. Soc. Bot. Jour. 27: 183. 1890. — An Hymeno- 
chaete pallida Cooke & Massee, Linn. Soc. Bot. Jour. 27: 97. 
1890? See Patouillard, Myc. Soc. Fr. Bui. 10: 78. 1894, and 
Burt, Ann. Mo. Bot. Gard. 5:367. 1918. 

Illustrations: Cooke, Grevillea 8: pi. 121^. f. 9; Australian 

Fungi, pi. ll.f.82. 

Type: in Kew Herb. 

Fructification coriaceous-papery, thin, pliant, resupinate and 

widely effused, sometimes reflexed, rarely umbonate sessile, 

the upper side tomentose, concen- 
trically sulcate, drying snuff-brown, 
weathering to cartridge-buff, the 
margin entire; hymenium even, vel- 
vety, snuff-brown to Benzo-brown; 

in structure 500-600 p. thick exclu- 
sive of the tomentose covering, com- 
posed of longitudinally and loosely 
interwoven, even-walled, pale-colored 
hyphae 3-3| n in diameter, which 

Fig. 32. S. papyrinum. Sec- g j ve tne } r co l or to the fructification, 




Ik 





J J Va 



tion of hymenium X 488, show- 
ing cystidia and paraphyses. 
From authentic specimen. 



the intermediate layer not dense on 
its upper side but grading into the 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 197 



tomentum; no conducting organs present; cystidia rather few 
and scattered, heavily and coarsely incrusted on the peripheral 



half, conical, 30-75 



i under the 



tation, confined to the hymenium; slender, flexuous paraphyses 
2\ n in diameter are abundant in the hymenium; spores hya- 
line, even, 4^-8X3-4 n — but few found. 

Resupinate on under side of limbs over areas up to 25X3? 
cm., and reflexed along both sides 1-2| cm. 

On under side of fallen limbs of frondose species. Florida, 
West Indies, Mexico, Colombia, and Brazil. October to May. 

Probably common. 

S. papyrinum belongs in the group with S. umbrinum and S. 
albo-badium; resupinate specimens of these species require ex- 
amination of sectional preparations for accurate determina- 
tion. The specimens which have been distributed by Ravenel 
and by Ellis in their exsiccati as S. papyrinum are S. umbrinum. 
In its reflexed stage, S. papyrinum is much more broadly reflexed 
than S. umbrinum and is concentrically sulcate; its cystidia are 
heavily incrusted and from 12 to 25 n in diameter by 30 to 75 ju 
long, while those of S. umbrinum are much longer in proportion 
to their diameter and often can be followed from deep in the 
subhymenium, taper so gradually and bear so little incrustation, 
and are so uniformly colored that some: mycologists have 
regarded them as setae, although they do not satisfy the defini- 
tion of setae. The cystidia of S. papyrinum are concolorous 
with the hyphae under the incrustation. S. albo-badium has 
cystidia heavily incrusted but smaller than those of S. papyrinum 
and not colored. 

On account of their structure, I have included in S. papyrinum 
the Cuban specimens listed by Berkeley & Curtis as S. mem- 
branaceum, for I find nothing to show that these specimens were 
ever compared with the type of the latter in Herb. Willdenow 
and collected on the Isle of Bourbon in the Indian Ocean; there 
is nothing in the original description of 8. membranaceum to 
show that this may not be more closely re'ated to S . fasciatum 
than to S. papyrinum. I have referred to S. papyrinum, as um- 
bonate-sessile forms, the specimen from Nicaragua distributed 
in Smith, Central Am. Fungi, 94, and a collection from Cuba by 
Underwood & Earle, 1584, which are cited below; these speci- 



i no ' Vo1 - 7 

ly» ANNALS OF THE MISSOURI BOTANICAL GARDEN 

mens have cystidia of the minimum dimensions given for the 
species and with less than the usual incrustation, as is the case 
with cystidia of the type of S. nicaraguense ; perhaps these two 
specimens are Hymenochaete pallida. 

Specimens examined: 
Exsiccati: Smith, Central Am. Fungi, 95 and 93 a and b. under 



Stereum rufo-fulvum (Mont.), and 94, und 



ame£ 



Florida: Adams Key, Dade Co., J. H. Small & C. A. Mosier, 

5364, comm. by N. Y. Bot. Card. Herb, (in Mo. Bot. Card. 

Herb., 71448); Miami, W. H. Long, 18310 (in Mo. Bot. 

Card. Herb., 55442); Palm Beach, R. Thaxtcr, 16 (in Mo. 

Bot. Gard. Herb., 43927). 
Cuba: type, from Montagne (in Kew Herb.), and C. Wright, 

274, and 240, both under the name S. membranaceum (both 

in Curtis Herb.); Alto Cedro, L. M. Underwood & F. S. 

Earle, 1481, 1492, 1584, comm. by N. Y. Bot. Gard. Herb.; 

Ceballos, C. J.Humphrey, 2726 (in Mo. Bot, Gard. Herb.); 

El Yunque Mt., Baracoa, L. M. Underwood & F. S. Earle, 

364 (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 

56714), and 739, 745, and 1233, comm. by N. Y. Bot. Gard. 

Herb.; .Managua, Earle & Murrill, 32, comm. by N. Y. 

Bot. Gard. Herb.; San Diego de los Bafios, Earle & Mur- 

64, 356, 362, 367, 380, all comm. by N. Y. Bot. Gard. 



Herb 



Porto Rico: Espinosa, J. A. Stevenson, 2751 (in Mo. Bot, Gard. 

Herb., 5554). 

Jamaica: A. E. Wight, comm. by W. G. Farlow; Hope Gardens, 

F. S. Earle, ltf, 165, 431, 494, all comm. by N. Y. Bot. 
Gard. Herb.; Port Maria, F. S. Earle, 467, comm. by N. Y. 
Bot. Gard. Herb.; Troy and Tyre, W. A. Murrill & W. 
Harris, 898, comm. by N. Y. Bot. Gard. Herb.; West- 
moreland, F. S. Earle, 425 A, comm. by N. Y. Bot. Gard. 
Herb.; San Juan, F. S. Earle, 62, comm. by N. Y. Bot. 
Gard. Herb. 

Mexico: Colima, W. A. & E. L. Murrill, 637, 648, comm. by 

N. Y. Bot. Gard. Herb, (in Mo. Bot, Gard. Herb., 54583, 
54584); Jalapa, C. L. Smith, in Smith, Central Am. Fungi, 
93a; Orizaba, W. A. & E. L. Murrill, 748, comm. by N. Y. 
Bot. Gard. Herb, (in Mo. Bot. Gard. Herb.. 54655). 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



199 



Nicara 



C. Wright, 264, type of S 



(in Curtis 



Herb.) ; Castillo Vieio, C. L. Smith, in Smith, Central Am 



Ometepe, C. L. Smith 



Smith. Central Am 



Fungi, 93b; San Juan del Norte, C. L. Smith, in Smith 
Central Am. Fund. 94. 



Canal 



Gatun, M. A. H. (in N. Y. Bot. Gard. Herb, and 



Mo. Bot. Gard. Herb., 56715). 
Colombia: Bonda, C.F. Baker, 26. 
Brazil: Santo Anna da Chapada, Matto Grosso, G. 0. Malme 

564, comm. by L. Romell. 



6i. S. Earlei Burt, n. sp. 



Plate 6, fig. 61. 



■HHBP 





Type: in Burt Herb, and N. Y. Bot. Gard. Herb. 

Fructification coriaceous-spongy, dry, effuso-reflexed, with 
the upper surface tomentose, snuff-brown, the margin entire; 
hymenium mouse-gray and some- 
what pruinose in the older portion, 
snuff-brown and veined toward the 
margin; in structure with the in- 
termediate layer 150 n thick, com- 
posed of longitudinally interwoven, 
colored hyphae 3-4 n in diameter, 
with the hymenial layer up to 200 
/* thick, zoned, containing cystidia 
in all its portions; cystidia colored, 
heavily hyaline incrusted on the _. „ „ , . a ,. , 

, ,f , , . Fig. 33. S. Earlei. Section of 

outer half, slender-pointed, 45-60 type y 6 8 ; cystidium, c, and spores, 
X5-12 n, protruding up to 30 n) *, x 4<S8. 

spores hyaline, even, 5-6X3-3^- 

Reflexed portion up to 1 cm. broad; resupinate portion lat- 
erally confluent for 8 cm., but a strip only 1 cm. wide removed 
from the substratum. 

In a wood pile. Hope Gardens, Jamaica. November. 

Fructifications of this species have the general aspect of those 
of S. papyrinum, but are thinner, more compactly inter- 
woven, with slenderer cystidia, and have the hymenial layer up 
to 200 m thick and composed of several zones; cystidia are 
present in each of these zones, and those of the innermost zones 
do not reach to the surface of the hymenium. In S. papyrinum 



200 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



the hymenium is a single layer of basidia, cystidia, and para- 
physes. In the collector's note, the color is given as "violet 
purple edged with white," but colors of dried specimens are as 
given above. 

Specimens examined: 
Jamaica: Hope Gardens, F. S. Earle, 151, type, comm. by N. Y. 

Bot. Gard. Herb. 



62. S. Chailletii Persoon, Myc. Eur. 1 : 125. 1822 (in 
****** Stereum of Thelephora); Fries, Epicr. 551. 1838; Hym. 
Eur. 642. 1874; Sacc. Syll. Fung. 6: 566. 1888; Bresadola, 



I. R. Accad. Agiati Atti III. 3: 106. 



1897. 



Plate 6, fig. 62. 



Thelephora Chailletii Pers. in Fries, Elenchus Fung. 1: 188. 



1828. 



Xerocarpus ambiguus Karsten, Soc. pro Fauna et Flora 



Fennica Actis a 1 : 38. 1881. — Trichocarpus ambiguus Karsten, 
Finska Vet.-Soc. Bidrag Natur och Folk 48: 407. 1889.— 
Hymenochaete ambigua Karsten in Sacc. Syll. Fung. 9: 230. 
1891. — Peniophora Atkinsonii Ellis & Everhart, Phila. Acad. 
Nat. Sci.Proc. 1894: 324. 1894; Sacc. Syll. Fung. 11: 129. 1895. 
Fructification coriaceous, nearly always resupinate, effused, 
occasionally reflexed, with upper surface tomentose, more or 

less concentrically sulcate when 
well developed, hair-brown to 
clove-brown, the margin entire; 
hymenium rather uneven 




' 



not 

polished, avellaneous to wood- 
brown; in structure 300-600 n 



thick, 



composed of somewhat 



longitudinally and not densely 
interwoven hyphae 3 



4 1 



/jl in 



Fig. 34. S. Chailletii. Section 
of hymenium X 666, showing para- 

physes; spores, 8. 



diameter, some of which are hya- 
line, thin-walled, and with deeply 
staining protoplasm, and many 
thick-walled, stiff, giving their 
color to the fructification, and 



curving into the hymenium where they terminate in cystidia; 
cystidia slightly colored, roughened above, 50-120 X 4-4.} /x, pro- 
truding up to 20 m, slender-pointed; spores white in spore col- 
lection, ellipsoidal, 5-6X3-3^- 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 201 



Wholly resupinate specimens |-2 cm, in diameter, becoming 
laterally confluent over areas up to 15X2 cm.; reflexed portions 
1-5 mm. broad — up to 2 cm. broad in European specimens. 

On dead Tsuga, Pseudotsuga, Abies, Picea, Larix, Thuja, 
and Cupressus. Canada to New Jersey, in Wisconsin, in Idaho 
to British Columbia and Washington, and in New Mexico at 
altitude 7500 ft. Occurs also in Europe. Probably throughout 
the year but most collections dated July to October. Infrequent. 
S. Chailletii occurs just often enough reflexed so that an 
observant collector will soon locate his gatherings correctly in 
Stereum. It is noteworthy by its colored cystidia of the same 
type as those of S. umbrinum but of only half the diameter 
of those of the latter, and by its occurrence on conifers of the 
species named above, and by restriction in geographic range to 
the northern United States and southern Canada and the Rocky 
Mountain plateau. The avellaneous, somewhat velvety hymen- 
ium is so uniform in appearance that when once learned this 
species may usually be recognized thereafter at sight. 

Specimens examined : 
Exsiccati: Ell. & Ev., N. Am. Fungi, 2904, under the name Hy- 

menochaete simulans Ell. & Ev., n. sp., but description does 

not seem to have been published; Kriegsr, Fungi Sax., 1202. 
Norway: Christiania, M. N. Blytt, determined by E. Fries (in 

Herb. Fries). 
Finland: Merimason, P. A. Karsten, authentic specimen of Tri- 

chocarpus ambiguus. 
Sweden: Stockholm, L. Romell, 24, 25, 341, all under the name 

Stereum abietinum. 
France: Arnac, Aveyron, A.Galzin, unnumbered spec, and 17948, 

comm. by H. Bourdot, 7926, and unnumbered respectively. 
Switzerland: Sachs, W. Krieger, in Krieger, Fungi Sax., 1202. 
Italy? or perhaps Hungary?: locality not given, G. Bresadola. 
Canada: Cow's Swamp, J. Macoun, 115; Dow's Swamp, J. 

Macoun, 249 in part. 
Ontario: Ottawa, J. Macoun, 57. 
Vermont: Ripton, E. A. Burt, two collections. 
New York: Beaver River, Adirondack Mts., G. F. Atkinson, 

Bot. Dept. of Cornell Univ., 4607; Ithaca, G. F. Atkinson, 

14189; Syracuse, G. F. Atkinson, 677, part of type of 

Peniophora Atkinsonii. 



202 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



New Jersey: Newfield, J. B. Ellis, in Ell. & Ev., N. Am. Fungi, 
2904. 

Wisconsin: M. C. Jensen, comm. by C. J. Humphrey, 2502 (in 

Mo. Bot. Gard. Herb., 5060). 
Idaho: Coolin, /. R. Weir, 11183, 11527, 11940 (in Mo. Bot. 

Gard. Herb., 56717, 56722, 56718); Kaniksu National 

Forest, Priest River, J. R. Weir, 65, 110 (the latter in Mo. 

Bot, Gard. Herb., 13272). 

British Columbia: Kootenai Mts., near Salmo, J. R. Weir, 482, 

510, 518 (in Mo. Bot. Gard. Herb., 18282, 3771, 1739); 

Sidney, J. Macoun, 81 (in Mo. Bot, Gard. Herb., 5887); 

Squamish, J. Macoun, 538 (in Mo. Bot. Gard. Herb., 55186). 
Washington: Bellingham, J. R. Weir, 7559 (in Mo. Bot, Gard. 

Herb., 55467, 55790); Stanwood, C. J. Humphrey, 7358 

(in Mo. Bot. Gard. Herb., 20103). 
New Mexico: Tejano Experiment Station, near Albuquerque, 

W. H. Long & P. W. Seay, comm. by W. H. Long, 21313 

(in Mo. Bot. Gard. Herb., 54884). 



63. S. ferreum Berk. & Curtis, Linn. Soc. Bot. Jour. 



10: 



332. 



1868; Sacc. Syll. Fung. 6: 586. 1888; Massee, Linn. 



Soc. Bot. Jour. 27: 197. 



1890. 



Plate 6, fig. 63. 



An Stereum areolatum Fries ? 

Type: in Kew Herb, and Curtis Herb. 

Fructifications corky, effused, usually resupinate, sometimes 
becoming barely reflexed on the upper side and there drab, 

nearly even; hymenium somewhat colliculose, 
not shining, cinnamon-drab to drab; in struc- 
ture up to 1 100 m thick, with the intermediate 
layer 500 n thick, bordered by a darker zone 
next to substratum and composed of colored, 
thick-walled, somewhat ascending, inter- 
woven hyphae 3—3^ n in diameter; hymenial 
layer up to 600 m thick, containing in all parts 

innumerable incrusted cystidia, minutely 

Fig. 35. S. ferreum. rough, either colored throughout or colored 

Section of hymenial under the i ncrustat ion, 20-25 X 5-7 




region of type, X 488. 



M, Pro- 



Shows rough/ colored trading up to 6 n; spores hyaline, even, 



cystidia. 



globose, 4 n in diameter, but few found. 



1920] 



BURT— THELEPHORACEAE OF NORTH AMERICA. XII 203 



Fructifications 4-8X1-2 cm., margin reflexed 1 mm. 

On bark of fibrous structure of an unrecorded species. Cuba 
and Jamaica. Rare. 

S. ferreum may be recognized by its resupinate, drab fructifi- 
cations, rarely having a narrowly pileate margin, and by the 
thick hymenial layer containing innumerable small colored 
cystidia which at the surface of the hymen ium have the colorless 
incrustation roughened. So few spores were observed that it 



may be the} 
related to S. 



S.f 



and I have been inclined to regard it as not specifically distinct 
from the latter, but we do not know yet that S. ferreum occurs 
on Taxus or a related genus; if not a strictly tropical species but 
a synonym of S. areolatum, the lack of a northern range in 



common 



eastern United States is at variance with species 
Europe and North America. 

Specimens examined: 
Cuba: C. Wright, 199, type (in Kew Herb.). 
Jamaica: Cinchona, W. A. & E. L. Murrill, 458, comm. by 

N. Y. Bot. Gard. Herb.; Sir John Peak, W. A. Murrill, 

803, comm. by N. Y. Bot. Gard. Herb. 



64. S. cinerascens (Schw.) Massee, Linn. Soc. Bot. Jour. 27: 
179. 1890. Plate 6, fig. 64. 

Thelephora cinerascens Schweinitz, Am. Phil. Soc. Trans. N. S. 
4: 167. 1832. — Hymenochaete cinerascens (Schw.) Leveille, Ann. 
Sci. Nat. Bot. III. 5: 152. 1846; Morgan, Cincinnati Soc. 
Nat. Hist. Jour. 10: 197. 1888. — Peniophcra cinerescens (Schw.) 
Sacc. in Sacc. Syll. Fung. 6: 646. 1888.— P. Schweinitzii 
Massee, Linn. Soc. Bot. Jour. 25: 145. 1889— Cor ticium as- 
chistum Berkeley & Curtis, Am. Acad. Arts & Sci. Proc. 4: 123. 
1858. — Peniophora Berkeleyi Cooke, Grevillea 8: 20. pi. 122. 



f. 4- 1879; Sacc. Syll. Fung. 6: 642. 1888; Massee, Linn. 
Soc. Bot. Jour. 25: 144. 1889. — Stereum moricola Berkeley, 
Grevillea 1: 162. 1873; Sacc. Syll. Fung. 6: 567. 1888. 
Peniophora moricola (Berk.) Massee, Linn. Soc. Bot. Jour. 
25: 141. 1889. — Stereum dissitum Berkeley, Grevillea 1: 164. 
1873. — Peniophora dissita (Berk.) Cooke, Grevillea 8: 150. 
1880; Sacc. Syll. Fung. 6: 645. 1888; Massee, Linn. Soc. 



204 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Bot. Jour. 25 



1889. — Corticium ephebium Berk. & Curt 



Grevill 



1: 178 



1873 



Sacc. Syll. Fung. 6: 618. 1888. 



Peniophora ephebia (Berk. & Curtis) Massee, Linn. Soc. Bot. 
Jour. 25: 151. 1889. — Stereum neglectum Peck, N. Y. State 
Mus. Rept. 33: 22. 1880. — Peniophora neglecta Peck, N. Y. 



State Mus 



40: 76 



& 



hart. Torr. Bot. Club 



277. 



1897; Sacc 




Fung. 



14: 224. 1900. 



loyd 



occidentalis (Ell. & Ev.) v. H 



& Litsch. K. Akad. Wiss. Wien Sitzungsber 



6: 791 



Stereum 



Lloyd, Myc. Writ. 4. Letter 



1907. 
5: 14. 



1914. 



Illustrations: Cooke, Grevillea 8: pi. 122. f. 4- 



1879. 



I 

1 






s 



Type: in Herb. Schweinitz, Curtis Herb., and Kew Herb. 
Fructifications coriaceous, often resupinate and effused, some- 
times reflexed, with upper surface strigose-hairy, concentrically 

sulcate, warm buff to pinkish buff, weathering 
gray, often laterally confluent, the margin 
entire; hymenium minutely bristly with the 
cystidia, even, drying pinkish buff to drab; in 
structure 400-600 n thick excluding the hairy 
covering, with the intermediate layer composed 
of longitudinally interwoven, thick-walled hy- 
phae 4-4 1 ^ in diameter; cystidia large, in- 
crusted, thick-walled, often brownish at the 
base, conical, 100-150 X 12-20 p, emerging up to 
40-70 n] spores white in spore collection, even, 
10-12X6 n y somewhat flattened on one side. 

Resupinate portions 1-10 X 1-2^ cm.; reflexed 
margin 2-8 mm. broad. 

On logs and fallen limbs of Ulmus, Tilia, 
Robinia, Morus, etc. Canada to Texas, west- 
ward to California, and in Mexico, Cuba, and 
Brazil. Common. June to February. 
Fully developed specimens of S. cinerascens may be recognized 
by their narrowly reflexed, strigose-hairy pileus and hymenium 
somewhat pruinose with the large, bristly, colorless cystidia. In 
sectional preparations, these cystidia are usually slightly colored 
at the base and more numerous and larger than in any other 
North American Stereum; the spores are very large also. 



Fig. 30. 

S. cinerascens. 

Cystidium, c, and 

spores, s, X 488. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 205 



Wholly resupinate specimens have merely a superficial resem- 
blance to Peniophora, for they are loosely attached to the 
substratum by the layer of loosely arranged, coarse hairs which 
forms the strigose covering of the upper surface of a reflexed 
specimen; the intermediate layer is well developed in resupinate 
specimens, and the cystidia and spores are the same as in reflexed 
specimens. It is surprising that a species so common and so 
marked in its microscopical characters should have seemed new 
so many times. 

Specimens examined: 
Exsiccati: Bartholomew, Fungi Col, 2337, 4648; Ell. & Ev., 

N. Am. Fungi, 2314, type distribution of Peniophora occi- 
dentalis; Shear, N. Y. Fungi, 313. 
Canada: J. Macoun, 1+5, 68, and another specimen comm. by 

J. B. Ellis, under the name Peniophora occidentalis ; Lower 
St. Lawrence valley, J. Macoun, 33, 34, 79. 
Quebec: Hull, J. Macoun, Nat. Hist. Surv. of Canada, 359, and 

J. Macoun, 52; Ironsides, /. Macoun, 282. 
Ontario: Guelph, J. H. Faull, Univ. Toronto Herb., 669 (in 

Mo. Bot. Gard. Herb., 44916); Jefferson, G. H. Graham, 
Univ. Toronto Herb., 673 (in Mo. Bot. Gard. Herb., 44922); 
Ottawa, J. Macoun, 234; Toronto. J. H. Faull, Univ. 
Toronto Herb., 651 (in Mo. Bot. Gard. Herb., 44947). 
Vermont: Middlebury, E. A. Burt, six collections. 
Massachusetts: W. G. Farlow, two collections. 
New York: Alcove, C. L. Shear, 1312, and in Shear, N. Y. Fungi, 

313; Cayuga Lake Basin, G. F. Aiizinson, 3020, 8023, J; 
Greenbush, C. H. Peck (in N. Y. State Mus. Herb, and 
Mo. Bot. Gard. Herb., 56020); Ithaca, C. J. Humphrey, 
261 , and a specimen comm. by G. F. Atkinson, Van Hook, 
comm. by G. F. Atkinson, 7988; Knowersville, C. H. Peck 
(in N. Y. State Mus. Herb, and Mo. Bot. Gard. Herb., 
55755) ; Syracuse, L. M. Underwood, 5 (in N. Y. Bot. Gard. 
Herb, and Mo. Bot. Gard. Herb., 56709); Verona, C. H. 
Peck, type of Stereum neglectum (in N. Y. State Mus. Herb., 
and perhaps a duplicate in Mo. Bot. Gard. Herb., 55754). 
Pennsylvania: Bethlehem, Schweinitz, type (in Herb. Schweinitz, 

Curtis Herb., and Kew Herb.). 
South Carolina: Curtis Herb., 5997, type of Stereum moricola 

(in Kew Herb.). 



onr IVoL - 7 

^!(Jb ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Georgia: Atlanta, E. Bartholomew, 569 ] 4 (in Mo. Bot. Gard. 

Herb., 44220), and in Bartholomew, Fungi Col., 4648. 
Florida: Cocoanut Grove, R. Thaxter, 95 (in Mo. Bot. Gard. 

Herb., 43922); Miami, W. H. Long, 12951 (in Mo. Bot. 

Gard. Herb., 55102); Totten Key, P. H. Rolfs. 
Alabama: Peters, 923, type of Corticium ephebium, 1004, 1007 

(in Curtis Herb., 6050, 6088, and 6089 respectively, and in 

Kew Herb.). 

Texas: C. Wright, Curtis Herb., 3903, type of Stereum dissitum 

(in Kew Herb., and probably a co-type in Burt Herb., and 
U. S. Dept. Agr. Herb.). 

Michigan: Ann Arbor, C. H. Kaufman, 25; New Richmond, 

C. H. Kauffman, 64 (in Mo. Bot. Gard. Herb., 19651). 

Ohio: Cincinnati, A. P. Morgan, comm. by Lloyd Herb., 2590, 

and A. P. & S. V. Morgan, comm. by U. S. Dept. Agr. 
Herb., under the name Hymenochaete imbricatula as deter- 
mined by Morgan; Linwood, C. G. Lloyd, 8553, 02835. 

Indiana: Hibernian Mills, Whetzel & Reddick, comm. by D. 

Reddick, 2. 

Minnesota: Cass Lake, J. R. Weir, 324 (in Mo. Bot. Gard. Herb., 
6968); Clearwater Lake, F. Weiss, 4 (in Mo. Bot. Gard. 

Herb., 56634); Wright Co., F. Weiss (in Overholts Herb., 

5367). 

Iowa: Webster, O. M. Oleson, 437 (in Mo. Bot. Gard. Herb., 

44060); Woodbine, C. J. Humphrey & C. W. Edgerton, 

comm. by C. J. Humphrey, 6535 (in Mo. Bot. Gard. Herb., 
14042). 

Missouri: Creve Coeur, P. Spaulding (in Mo. Bot. Gard. Herb., 

5137); Upper Creve Coeur, E. A. Burt (in Mo. Bot. Gard. 
Herb., 56711). 

Arkansas: Fordyce, C. J.Humphrey, 5778. 

Nebraska: Lincoln, C. L. Shear, 1052; Pawnee City, C. L. Shear, 

1016. 

Kansas: Louisville, E. Bartholomew, in Bartholomew, Fungi 

Col., 2337; Rooks Co., E. Bartholomew (in Burt Herb, and 
Mo. Bot. Gard. Herb., 5011). 

Montana: F. W. Anderson, in Ell. & Ev., N. Am. Fungi, 2314. 

California: Bear Valley, near Olema, M. A. H. (in N. Y. Bot. 

Gard. Herb, and Mo. Bot. Gard. Herb., 56591); Berkeley, 



1920] 



BURT— THELEPHORACEAE OF NORTH AMERICA. XII 



207 



H. A. Lee, comm. by W. A. Setchell 
Gard. Herb.. 44304). 



(in Mo 



Mexico: Xuchiles, near Cordoba, W. A. & E. L. Murrill 



Herb 



comm. by N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard 



Nicaragua: C. Wright, 27 L 



of Cortici 



aschistum and 



Peniophora Berkeleyi (in Curtis Herb 



Cuba: C. G. Lloyd, 428 (in Mo. Bot. Gard. Herb., 55157); Alto 

Cedro, Earle & Murrill, 515 (in N. Y. Bot. Gard. Herb, and 
Mo. Bot. Gard. Herb., 56291); Havana, Bro. Leon, comm. 
by J. R. Weir, 10188 (in Mo. Bot. Gard. Herb., 56216). 

Jamaica: Chester Vale, W. A. & E. L. Murrill, 343, comm. by 

N. Y. Bot. Gard. Herb. 

Brazil: Matto Grosso, Santa Anna da Chapada, G. V. Malme, 

572, comm. by L. Romell. 



65. S. magnisporum 

Type: in Burt Herb. 



ng 



Fructifications coriaceous-gelatinous, thin, resupinate, becom 
ing confluent, free all around, with margin reflsxed on the uppe] 
side, probably white, drying pale pinkish 
buff, hoary, the margin white, entire; hy- 
menium even or with one or two broad 
veins, setulose with the large cystidia, 
drying pinkish buff; in structure 300 /u 
thick when dry, swelling to 1200-1500 
n thick when wet for sectioning, of gel- 
atinous consistency, composed of loosely 
interwoven, hyaline hyphae 2 
diameter, not incrusted; hymenial layer 
not zonate, composed of large simple 
basidia 45-60X15 ft, having 4 sterig- 
mata 12 M long, of hyaline, filiform, 
flexuous paraphyses 2-2£ y. in diameter, 



n in 



exceeding the basidia, and of 




incrusted cvstidia 



90X12-15 



M 



protruding up to 60 m; spores hyal 
even, 15-20 X 12-14 u. 



) 



Fig. 37. S. magnisporum. 
Cystidium, c, basidia, 6, and 
spores, s, X 488. From type. 



Fructifications 2-6 mm. in diameter, laterally confluent for 



mm,, margin reflexed 



mm 



[Vol. 7 



208 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



On dead limbs of a frondose species. Jamaica. December 

to January. 

S. magnisporum may be recognized by its small, whitish 
fructifications, with narrowly reflexed or free margin, pale hy- 
menium distinctly setulose with the large cystidia, and by the 
very large spores. The large spores and basidia show relation 
of S. magnisporum to Aleurodiscus, but the absence of granular 
matter or of any unusual character of the paraphyses leads to 
the belief that this species will usually be sought for among the 

Stereums. 

Specimens examined: 
Jamaica: Chester ValeJ W. A. & E. L. Murrill, 328, type, coram. 

by N. Y. Bot. Gard. Herb.; Cinchona, W. A. & E. L. 



Murrill, 522, comm. by N. Y. Bot. Gard. Herb. 



66. S. spumeum 



fi 



Corticium spumeum Berk. & Rav. in Curtis Herb, (in part); 
Grevillea 20: 13. 1891 (in part— nomen).— C. ochroleucum, "as 
resupinate ambient condition," Berk. & Curtis, Grevillea 1: 
166. 1873, but not Stereum ochroleucum Fries.— Not Corticium 
ochroleucum var. erimosum Berk. & Curtis, Grevillea 1: 166. 

1873. 

Type: in Burt Herb. 

Fructifications spongy-soft, effused, resupinate, separable, 
sometimes narrowly reflexed, the upper surface tomentose and 
becoming cartridge-buff to pinkish buff in the herbarium, the 
margin entire; in structure 400-1500 m thick, composed of 
loosely interwoven, hyaline, thick-walled hyphae 3-4§ m in di- 
ameter, sometimes nodose-septate, the intermediate layer not 
bordered on its upper side by a crust-like or colored zone; 
hymenium even, cream-buff to pinkish buff; no conducting 
organs; cystidia incrusted, 36-60X9-12 n, sometimes protrud- 
ing up to 40 m; spores hyaline, even, 5-9X3-4 m. 

Resupinate over areas 1-10 X 1-5 cm., reflexed portion 1-4 

mm. broad when present. 

On bark and wood of dead beech, oak, and other frondose 
limbs. New York to Mexico. August to January. Rare. 

S. spumeum is noteworthy by its narrowly reflexed pileus, 
soomrv-soft throughout, and without differentiation of its sur- 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 209 



face of soft, matted, interwoven hairs from i;he hyphae of the in- 
termediate region, by its buff hymenium, and by its incrusted 
cystidia. These incrusted cystidia and different aspect of the 
fructifications afford sharp separation from S. ochraceo-ftavum ; 
S. ochroleucum and S. rugosiusculum have the general aspect of 
S. spumeum but both lack incrusted cystidia, and S. rugosius- 
culum has in its subhymenial region pyriform, vesicular organs. 
S. spumeum is so frequently resupinate or very narrowly reflexed 
that gatherings are likely to be referred to Peniophora. 

Specimens examined: 
New York: Hudson Falls, S. H. Burnham, 27 (in Mo. Bot. 

Gard. Herb., 54486). 

Pennsylvania: E. Michener, 186 '4 (in Curtis Herb., under the 

name Corticium giganteum). 
South Carolina: Aiken, on oak limbs, H. W. Ravenel, 1772 (in 

Curtis Herb., under the name Corticium ochroleucum, 



u 



formerly C. spumeum"). 



Louisiana: Baton Rouge, Edgerton & Humphrey; St. Martin- 

ville, A. B. Langlois,E, type 
Mexico: Guernavaca, W. A. & E. L. Murrill, 405, 413, 414, 498, 

503, 520, comm. by N. Y. Bot. Gard. Herb, (in Mo. Bot. 

Gard. Herb., 54520-54523, 56685, 55524); Cordoba, W. A. 

& E. L. Murrill, 1214, comm. by N. Y. Bot. Gard. Herb. 

(in Mo. Bot. Gard. Herb., 54592). 



67. S. erumpens Burt, n. sp. Plate 6, fig. 67. 

Type: in Burt Herb. 

Fructifications corky, rarely resupinate, usually bursting out 
from the inner bark as small pezizaeform, orbicular disks or cups 
with elevated black margins and cinereous or pallid neutral gray 
hymenium; these fructifications may become crowded as if 
confluent, and then broken up into frus- 
tules and remain attached by the under 



side to the substratum, or the margin on 

the upper side may grow outward so as 

to form umbonate, sessile pilei attached 

by the umbo and lower side, with the 

upper surface narrowly concentrically Fig 38 s enmpm8 

sulcate, mummy-brown to fuscous; hy- Section of type, x 90. 

9 




|Vol. 7 
210 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



menium even or somewhat tubercular, pallid neutral gray; 
in structure 200-300 n thick, composed of ascending, densely- 
interwoven hyphae both colored and hyaline, the former 3| m in 
diameter, with the tips arranged side by side in colored subhy- 
menial zones, mark the 1-3 strata finally present; cystidia 
incrusted, cylindric, 30-60 X 8-20 n, sometimes protruding up to 

20 m beyond the hymenium, starting from all parts of the fructi- 
fication; spores hyaline, even, 5-7Xl§-2§ n. 

Fructifications 1-2| mm. in diameter, reflexed 1-2 mm. 
On dead limbs of alder, chestnut, willow, and other frondose 
species. Rhode Island to Alabama and westward to Washing- 
ton and Oregon. March to January. Occasional. 

S. erumpens combines the characters of S. versiforme and 
Peniophora cinerea; it is more constantly and distinctly reflexed 
than S. versiforme, always has a gray hymenium, and has quite a 
different mode of origin from the latter. In the type small 
blackish bodies burst out from the bark, open at the tip, dis- 
closing whitish hymenium, and then grow to mature condition. 
Specimens at hand do not show how such a large resupinate 
fructification as that collected by E. T. and S. A. Harper, No. 
819, cited below, does arise, and I may be wrong in referring the 
specimen to S. erumpens. An important microscopical detail of 
S. erumpens is the narrow olivaceous zone of colored hyphal tips 
at the very base of the basidia of the hymenium. 

Specimens examined: 
Exsiccati: Ellis, N. Am. Fungi, 720, under the name Corticium 

quercinum var. scutellatum. 
Rhode Island: Lincoln, F. W. Collins. 
New York: East Galway, E. A. Burt; Ithaca, C. J. Humphrey, 

2568 (in Mo. Bot. Gard. Herb., 20784); Karner, H. D. 

House (in N. Y. State Mus. Herb, and Mo. Bot. Gard. 

Herb., 55210); New Scotland, C. H. Peck (in N. Y. State 

Mus. Herb., T 28, and Mo. Bot. Gard. Herb., 54658). 
New Jersey: Newfield, J. B. Ellis, in Ellis, N. Am. Fungi, 720. 
Maryland: Takoma Park, C. L. Shear, 959. 
District of Columbia: North Takoma, C. L. Shear, 1043, type. 
Georgia: Raleigh, R. M. Harper, 2087b, comm. by P. L. Ricker, 

and (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 

42597). 



1920 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 211 



Alabama: Auburn, F. S. Earle, 2301 (in N. Y. Bot. Gard. Herb. 

and Mo. Bot. Gard. Herb., 56292). 
Indiana: Scottsburg, J. R. Weir, 5886 (in Mo. Bot. Gard. Herb.> 

55462). 
Illinois: Glencoe, E. T. & S. A. Harper, 819, 987. 
Arkansas: Fayetteville, R. R. Rosen, comm. by L. 0. Overholts, 

5117 (in Mo. Bot. Gard. Herb., 56358). 
Montana: Missoula, J. R. Weir, 854 (in Mo. Bot. Gard. Herb.,, 

9435). 
Washington: Brewerton, E. Bartholomew, comm. by N. Y. Bot- 

Gard. Herb, (in Mo. Bot. Gard. Herb., 4939). 
Oregon: Grants Pass, J. R. Weir, 8701 (in Mo. Bot. Gard. Herb., 

36742). 



68. S. sulcatum Burt in Peck, N. Y. State Mus. Rept 
154. 1901; Lloyd, Myc. Writ. 5. Notes 44: 619. text f. 



1917. 



Lloyd 



fig. 68 



Type: in Burt Herb., N. Y. State Mus. Herb., and Bresadola 
Herb. 

Fructification corky, rigid, resupinate or effuso-reflexed, with 
the reflexed part becoming glabrous, bister, irregular, deeply and 
concentrically sulcate; hymenium uneven or somewhat tuber- 
cular, not polished, drying between light buff and pinkish buff, 
assuming a reddish color where bruised; in structure 600-1500 11 
thick, with the intermediate layer 
bordered by a dark dense zone on its 
upper side, and composed of very 
densely and longitudinally inter- 
woven, hyaline hyphae 3-3§ /* in 
diameter, the hymenial layer be- 
coming zonate or stratose; no col- 
ored conducting organs; cystidia 
incrusted, 30-50X8-12 **; spores 
white in spore collection, even, sub- 
globose, 4-6X3-5 /x. 

Confluent over areas 3-15X1-8 

n t . n 1A Fig. 39. S. sulcatum. Section 

cm.; reflexed margin 3-10 mm. of hymen ial region x 90; cystidia, 




broad. 



c, and spores, s, X 665. 



[Vol. 7 
212 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



On logs and stumps of Tsuga, Abies, Picea, Taxodium, Pseu- 
dotsuga, and Larix. Canada to Texas and westward to British 
Columbia and Washington. May to November. Frequent. 

S. sulcatum may be recognized by its brown, deeply and sharp- 
ly and concentrically sulcate pileus, ruddy hymenium, incrusted 
cystidia, and occurrence on conifers. Where the northern 
hemlock occurs it is usually on this species. S. Chailletii is 
found on conifers throughout the same northern geographical 
range, but is much thinner and does not have as large nor 
incrusted cystidia. In the older herbaria S. sulcatum is often 
found under the name Stereum rugosum, to which specimens 
were erroneously referred. 

Specimens examined: 
Exsiccati: Ell. & Ev., N. Am. Fungi, 1935, under the name 

Stereum rugosum; Ell. & Ev., Fungi Col., 217, under the 

name S. rugosum. 
Canada: J. Macoun, 27, 32, 43; Lower St. Lawrence Valley, 

J. Macoun, 69a, 76. 

Ontario: Ottawa, J. Macoun, 234, and in Ell. & Ev., N. Am. 

Fungi, 1935. 

New Hampshire: North Conway, L. 0. Overholts & H. H. York, 

comm. by L. 0. Overholts, 5033 (in Mo. Bot. Gard. Herb., 
56350). 

New York: Floodwood, E. A. Burt, type; Ithaca, G. F. Atkin- 
son, 2023, 2617, 2636, 5072, 7889, 19398, and C. 0. Smith, 
comm. by G. F. Atkinson, 8032; North Elba, C. H. Kauff- 
man, 7 (in Mo. Bot. Gard. Herb., 21821); Pompey, L. M. 
Underwood, in Ell. & Ev., Fungi Col., 217. 

Louisiana: Lutcher, H. von Schrenk, 26 (in Mo. Bot. Gard. 

Herb., 42637). 

Texas: Houston, H. W. Ravenel, 113 (in U. S. Dept. Agr. Herb., 

under the herbarium name Stereum tricolor). 

Wisconsin: Ladysmith, C. J. Humphrey, 1908 (in Mo. Bot. 

Gard. Herb., 42917). 

West Virginia: comm. by W. G. Farlow. 

Tennessee: Elkmont, C. H. Kauffman, 60 (in Mo. Bot. Gard. 

Herb., 16403). 

Montana: Gallatin National Forest, Spring Hill, G. G. Hedgcock t 

comm. by C. J. Humphrey, 2164 (in Mo. Bot. Gard. Herb., 
10399). 



1920 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



213 



Idaho: Kaniksu National Forest, Priest River, J. R. Weir, 4, 

29, 58, 74, 82, and 102 (the last in Mo. Bot. Gard. Herb., 

16029). 
Canadian Rocky Mts.: Lake Louise, J. Macoun, 3; Lake 

O'Hara, J. Macoun, 7 ; Papiston Creek, J. Macoun, 8. 
British Columbia: Yoho Valley, J. Macoun, 5. 
Washington: Mt. Paddo, W. N. Suksdorf, 843, 844. 
Oregon: Sumpter, G. G. Hedgcock, comm. by C. J. Humphrey, 

2570 (in Mo. Bot. Gard. Herb., 20460). 



69. S. subpileatum Berk. & Curtis, Hooker's Jour. Bot. 1: 
238. 1849; Grevillea 1: 163. 1873; Sacc. Syll. Fung. 6: 585. 
1888; Massee, Linn. Soc. Bot. Jour. 27: 192. 1890; Long, Jour. 



1915. 



Plate 6, fig. 69. 



Agr. Res. 5: 421. pi. 41- 

Illustrations: Jour. Agr. Res. 5: pi. 41- 

Type: in Curtis Herb, and Kew Herb. 

Fructifications thick, corky, drying rigid, very hard, resupi- 
nate oreffuso-reflexed, sometimes laterally confluent and attached 
by the umbos, with upper surface concentrically sulcate, some- 
what zonate, tomentose, cinnamon-brown, the margin entire; 
hymenium even, light buff; in structure 800-1200 m thick, with 
the intermediate layer bordered and 
connected with the tomentum by a 
denser and darker crust and bearing 
on the opposite side a hymenial 
layer which becomes multizonate; 
hyphae of intermediate layer col- 
ored, thick-walled, stiff, 3-3f m in 
diameter, densely and longitudi- 
nally arranged; cystidia incrusted, 



cylindric, 30-36X7 m, becoming col- 
ored where buried in older zones of 
the hymenium, at first sometimes 



slightly aculeate; 






spores hyaline, 
even, 4-5X3 ». 

Fructifications with reflexed por- 
tion 1-6 cm. broad. 

Perennial on logs of several 




Fig. 40. S. subpileatum. Sec- 
tion X 68; hymenium, h } crust-like 



Species of QuerCUS Causing a pock- zone, z, cystidia of type, c, X 488. 



[ Vol. 7 



214 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



eted or honeycomb heart rot. North Carolina and Ohio to 

Mexico, and in Cuba. 

In general aspect S. subpileatum is not distinguishable from 
S. sepium and S. insigne; it is more commonly met with than 
these latter species and with them occurs on oak logs, is also 
tobacco-colored and sulcate above and has a whitish hymenium 
which differs from the other species of this group by containing 
cylindric, incrusted cystidia and only very rarely an occasional 
paraphysis with its outer portion of bottle-brush or aculeate 
form. Usually such paraphyses are not found in preparations 
of the hymenium of this species. Occasionally preparations 
may show young cystidia which are merely rough above or some- 
what aculeate. One must not confuse S. subpileatum with the 
other species which have numerous and conspicuous bottle- 
brush paraphyses. 

Specimens examined: 
Exsiccati: Ell. & Ev., Fungi Col., 917; Ravenel, Fungi Am., 219; 

Ravenel, Fungi Car. i: 30; Smith, Cent. Am. Fungi, 140. 
North Carolina: Blowing Rock, G. F. Atkinson, J+183. 
South Carolina: Santee, H. W. Ravenel, type (in Curtis Herb., 

1007); Society Hill (in Curtis Herb., 1002). 
Georgia: Vienna, C. J.Humphrey, 5228. 
Florida: W. W. Calkins (in U. S. Dept. Agr. Herb., Burt Herb., 

N. Y. Bot. Gard. Herb., and Mo. Bot. Gard. Herb., 56759), 

and in Ell. & Ev., Fungi Col., 917. 
Alabama: Auburn, F. S. Earle & C. F. Baker (in Burt Herb, and 

Mo. Bot. Gard. Herb., 5110); Montgomery Co., R. P. 

Burke, 31 (in Mo. Bot. Gard. Herb., 17137). 
Louisiana: St. Martinville, A. B. Langlois. 
Ohio: A. P. Morgan (in Lloyd Herb., 2607). 
Kentucky: Mammoth Cave, C. G. Lloyd, 2798. 
Missouri: Columbia, B. M. Duggar, 550; Marianna, H. von 

Schrenk (in Burt Herb, and Mo. Bot. Gard. Herb., 42837); 

Wicks, L. O. Overholts, 3161 (in Mo. Bot. Gard. Herb., 

5713). 
Arkansas: W. H. Long, 12703, 18502 (in Mo. Bot. Gard. Herb., 

44160, 44161). 
Texas: Jasper, E. R. Hodson, 325, comm. by P. L. Ricker. 
Mexico: Jalapa, C. L. Smith, in Smith, Cent. Am. Fungi, 146. 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



215 



Cuba: C. Wright, 515, the S. scytale of Fungi Cubenses but 

according to the type (in Curtis Herb.). 



70. S. sepium Burt, n. sp. Plate 6, fig. 70. 

Type in Burt Herb. 

Fructification corky, drying rigid, hard, resupinate, becoming 
broadly reflexed, with the upper surface concentrically sulcate, 
somewhat zonate, tomentose, sepia, 
the margin paler and entire; hyme 
nium even, not shining, between ligh 
buff ai 
600-151 



d avellaneous; in structure 



thick 



mm 



in resupinate portion of M 



specimens 



> 



with the intermediate 



bordered and connected with 



the tomentum by a denser 
darker zone and bearing on th 
posite side a hymenial layer 1 
becomes multizonate; hypha 



d 



of 





termediat 



layer 



d. thick- 



ailed, densely and 



ar- 



d 



diameter; cystidia 



Fig. 41. S. sepium. Hyme- 
nium of type X 665, showing 
cystidia, c, and bottle-brush 
paraphyses, p. 



incrusted, cvlindric 



35X7 ix, be 



coming colored where buried in the deeper zones of the hyme- 
nium; paraphyses of bottle-brush or aculeate form, numerous and 
conspicuous in the hymenial surface, cylindric, 12-25X3-5 n) 

spores hyaline, even, 4X2|ju. 

Probably resupinate over large areas, for fragments fractured 



des are 6 cm. sauare: reflexed mar 



cm 



6 cm 



Under side of rotten logs of frondo 



Pennsylvania 



to Mexico and Colombia. Collected from July to December 

but probably perennial. 

The few collections of S. sepium which have been observed 
have the upper surface of the pileus a little brighter colored than 
that of S. subpileatum and the hymenium more avellaneous, but 



I cannot certainly 



the former from 



by the very numerous and conspicuous bottle-brush paraphyses 
which are present, in addition to cystidia, in the hymenium of 



o i r |VoL - 7 

21b ANNALS OF THE MISSOURI BOTANICAL GARDEN 



S. sepium. The specimens of Mexican collections cited below 
have larger size than those from the United States. 
Specimens examined: 

Exsiccati: Ellis, N. Am. Fungi, 1205, under the name Stereum 

subpileatum. 

Pennsylvania: West Chester, Everhart & Haines, in Ellis, N. 

Am. Fungi, 1205. 

North Carolina: Blowing Rock, G. F. Atkinson. 

South Carolina: Clemson College, P. H. Rolfs, 1632. 

Georgia: Vienna, C. J. Humphrey, 5229, type. 

Mexico: Jalapa, W. A. & E. L. Murrill, 117, 188, comm. by 

N. Y. Bot. Gard. Herb, (in Mo. Bot, Card. Herb., 11011, 
54445), and 39 (in N. Y. Bot, Gard. Herb, and Mo. Bot. 
Gard. Herb., 56760). 

Colombia: Bonda, C. F. Baker, 24, in Plants of Santa Marta, 

Colombia, under the name Stereum illudens. 



71. S. albobadium (Schw.) Fries, Epicr. 551. 1838; Mor- 
gan, Cincinnati Soc. Nat. Hist. Jour, io: 195. 1888; Sacc. Syll. 
Fung. 6: 579. 1888; Massee, Linn. Soc. Bot. Jour. 27: 194. 

Plate 6, fig. 71. 



1890. 



Thelephora albobadia Schweinitz, Naturforsch. Ges. Leipzig 
Schrift. 1: 108. 1822 (in C. Corticia); Am. Phil. Soc. Trans. 
N. S. 4:167. 1832; Fries, Elenchus Fung. 1: 189. 1828. 
T. albo-marginata Schweinitz in Berkeley, Hooker's London 
Jour. Bot. 6: 324. 1847; Lea's Cat. Plants Cincinnati, 66. 
1849; Sacc. Syll. Fung. 6: 539. 1888.— Peniophora albo- 
marginata (Schw.) Massee, Linn. Soc. Bot, Jour. 25: 144. 
1889. — Stereum bizonatum Berkeley & Curtis, Grevillea 1 : 163. 
1873; Sacc. Syll. Fung. 6: 582. 1888; Massee, Linn. Soc. Bot. 
Jour. 27: 178. 1890.— S. Coffearum Berkeley & Curtis, Linn. 
Soc. Bot. Jour. 10: 332. 1868; Sacc. Syll. Fung. 6: 576. 1888. 

Hymenochaete paupercula Berkeley & Curtis, Linn. Soc. Bot. 
Jour. 10: 334. 1868.— Peniophora paupercula (Berk. & Curtis) 
Cooke, Grevillea 8: 150. 1880; Sacc. Syll. Fung. 6: 645. 1888. 
Type: I was unable to find the type in Herb. Schweinitz, 
although it was studied by Berkeley & Curtis, Acad. Nat. Sci. 
Phila. Jour. 3: 221. 1856. 

Fructifications coriaceous, thin, at first resupinate, orbicular, 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



217 



S 










becoming confluent, sometimes becoming narrowly reflexed, 
with the upper surface villose, varying from bufify brown to 
Natal-brown, becoming somewhat zonate when reflexed about 
5 mm., the margin entire and 
usually whitish; hymenium even, 
somewhat velvety, bister or snuff- 
brown, becoming light drab and 
somewhat pruinose with age; in 
structure about 500 \x thick, the 
intermediate layer with a darker 
zone on its upper side and com- 
posed of loosely, longitudinally 
arranged, slightly colored hyphae 
3-3| ft in diameter; hymenium 
30-45 ft thick, not zonate, having 

incrusted cystidia 30-45X8-15 n all confined to the single- 
layered hymenium, protruding up to 25 n; branched, filiform 
paraphyses 2 m in diameter, becoming colored, are present also 
in the hymenium, basidia simple, 4-spored; spores white in 
spore collection, even, flattened on one side, 6-11 X3-4§ m. 

Fructifications 5-10 mm. in diameter, becoming confluent 
over areas 1-2 cm. wide and 3 to many cm. long, and reflexed 

2-5 mm. 

On dead frondose wood and fallen limbs. New York to 
Mexico and westward to Idaho and Arizona, in the West Indies, 



Fig. 42. S. albobadium. Section 
X 90; cystidium, c, paraphyses, p, 
and spores, s, X 665. 



d reported from 



hout the 



Common 



S. albobadium may usually be recognized by its brown, velvety 
hymenium with a white border; with age the hymenium tends 
to become more uniformly light drab or pruinose, but some 
small fructifications in the vicinity are likely to show the original 



color contrasts. 



has a wide geographic range and 



somewhat variable in coloration but is verv constant in micro 



the branched, colored 



highly 



distinctive 



Specimens examined: 



Exsiccati: Bartholomew, Fungi Col., 3688, 4784; Ellis, N. Am. 
. Fungi, 15; Ravenel, Fungi Am., 221, 449; Ravenel, Fungi 
Car. i : 29. 
New York: Grand View, H. von Schrenk (in Mo. Bot. Gard. 



[Vol. 7 
218 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Herb., 43009) ; Orient, R. Latham (in Mo. Bot. Gard. Herb., 
16267). 

New Jersey: Newfield, J. B. Ellis, in Ellis, N. Am. Fungi, 15. 

Maryland: Plummers Island, C. L. Shear, 1276, 1277; Seven 

Locks, P. L. Richer, 1007; Takoma Park, C. L. Shear, 1118, 

1126. 

District of Columbia: Washington, C. L. Shear, 1263-1265, 1402. 
Virginia: Arlington Cemetery, W. H. Long, 12978 (in Mo. Bot. 

Gard. Herb., 55104). 

North Carolina: Chapel Hill, W. C. Coker, 3849 (in Mo. Bot. 

Gard. Herb., 56672). 

South Carolina: Curtis Herb., 1924, type of Stereum bizonatum 

(in Kew Herb.); Ravenel, in Ravenel, Fungi Car. i: 29; 
Aiken, H. W. Ravenel, in Ravenel, Fungi Am., 449; Clem- 
son College, P. H. Rolfs, 1637; Society Hill, under the 
name T. albo-mar ginata (in Curtis Herb.). 

Georgia: Atlanta, E. Bartholomew, in Bartholomew, Fungi Col., 

4784; Darien, H. W. Ravenel, in Ravenel, Fungi Am., 221. 

Florida: New Smyrna, C. G. Lloyd, 2089, 2104, 2132. 

Alabama: Auburn, F. S. Earle (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56764), F. S. Earle & C. F. Baker 
(in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 5055, 
56765, 56772), C. R. Hudson (in N. Y. Bot. Gard. Herb, 
and Mo. Bot. Gard. Herb., 55568); McGeher (in N. Y. Bot. 
Gard. Herb, and Mo. Bot. Gard. Herb., 56766), and L. M. 
Underwood, comm. by U. S. Dept. Agr.; Fayette Co., 
P. V. Siggers, comm. by A. H. W. Povah, 16 (in Mo. Bot. 
Gard. Herb., 14849); Mobile, E. Bartholomew, 5752 (in 
Mo. Bot. Gard. Herb., 44257) ; Montgomery, R. P. Burke, 
5, 29 (in Mo. Bot. Gard. Herb., 20914, 17071). 

Mississippi: Ocean Springs, F. S. Earle, 181 (in Mo. Bot. Gard. 

Herb., 44311). 

Louisiana: St. Martinville, A. B. Langlois. 

Texas: Paris, C. L. Shear, 1234; Quitman, W. H. Long, 18448, 

12081 (in Mo. Bot. Gard. Herb., 55105,55131); San Anto- 
nio, H. von Schrenk, also W. H. Long, 21217 (in Mo. Bot. 
Gard. Herb., 42577 and 55131 respectively). 

Ohio: C. G. Lloyd, 189, 594 (in Lloyd Herb.); College Hill, C. 

G. Lloyd, P; Norwood, C. G. Lloyd, 2810. 






19201 o 1 n 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII Zl\) 



Missouri: Meramec, P. Spaulding (in Mo. Bot. Gard. Herb., 

5017) ; Perry ville, L. 0. Overholts & R. A. Studhalter, comm. 
by L. 0. Overholts, 2723 (in Mo. Bot. Gard. Herb., 44293) ; 
Upper Creve Coeur, E. A. Burt (in Mo. Bot. Gard. Herb., 

54861, 56768). 
Kansas: Rooks Co., E. Bartholomew (in Burt Herb, and Mo. 

Bot. Gard. Herb., 5054); Stockton, E. Bartholomew, in 

Bartholomew, Fungi Col., 3688. 
Idaho: Bonner's Ferry, J. R. Weir, 592 (in Mo. Bot. Gard. 

Herb., 36746). 
Arizona: Phoenix, W. H. Long, 19030 (in Mo. Bot. Gard. Herb., 

55106). 

New Mexico: Cienega Springs, W. H. Long, 21525 (in Mo. Bot. 

Gard. Herb., 55155); Tyom Experiment Station, W. H. 
Long, 21364, 211+08 (in Mo. Bot. Gard. Herb., 55107, 55108) ; 
Tejano Experiment Station, W. H. Long, 21889, 21897, 
21902 (in Mo. Bot. Gard. Herb., 55165-55167). 

Bermuda: S. Brown, N. L. Britton, & F. J. Seaver, 1244 (in 

N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. Herb., 56273). 

Cuba: C. Wright, 247, type of Stereum Coffearum (in Curtis 

Herb.), and 542, type of Hymenochaete pauper cula (in 
Curtis Herb.), and C. G. Lloyd, 423 (in Mo. Bot. Gard. 
Herb., 55159); Alto Cedro, L. M. Underwood & F. S. 
Earle, 1492, 1590, comm. by N. Y. Bot. Gard. Herb.; 
La Gloria, Camaguey, J. A. Shafer, 740 (in N. Y. Bot. 
Gard. Herb, and Mo. Bot. Gard. Herb., 56770) ; Managua, 
Earle & Murrill, 11, comm. by N. Y. Bot. Gard. Herb.; 
Omaja, C. J. Humphrey, 2746 (in Mo. Bot. Gard. Herb., 
14385) ; San Diego de los Bafios, Earle & Murrill, 281 , 302, 
316, 353, comm. by N. Y. Bot. Gard. Herb. 

Porto Rico: Rio Piedras, J. A. Stevenson, 2424, 6272 (in Mo. 

Bot. Gard. Herb., 3607, 55090). 

Mexico: Jalapa, W. A. & E. L. Murrill, 301, 309, comm. by 

N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard. Herb., 54432, 
54483); Motzorongo, Cordoba, W. A. & E. L. Murrill, 992, 
comm. by N. Y. Bot. Gard. Herb, (in Mo. Bot. Gard. 
Herb., 54597); Orizaba, W. A. & E. L. Murrill, 760, 761, 
766, 769, 774, 779, comm. by N. Y. Bot. Gard. Herb, (in 
Mo. Bot. Gard. Herb., 54627, 54631, 54628, 54629, 54610, 



220 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



(Vol. 7 



54645); Tepeite Valley, Guernavaca, W. A. & E. L. 
Murrill, 408, comm. by N. Y. Bot. Gard. Herb, (in Mo. 
Bot. Gard. Herb., 54544); Xuchiles, Cordoba, W. A. & 
E. L. Murrill, 1209, 1210, comm. by N. Y. Bot. Gard. Herb. 
(in Mo. Bot. Gard. Herb., 54598, 54599). 



7 



S. heterosDorum 



n. sp 



Plate 6, fig. 72 



Mo. Bot. Gard. Herb 



Fructifications coriaceous, thin, resupinate, orbicular, becom- 
ing confluent, sometimes reflexed, with the upper surface villose, 
bister, somewhat concentrically sulcate and zonate, the margin 
entire, whitish ; hymenium even, somewhat velvety, bister, becom- 
ing light drab and somewhat pruinose in the center with age; in 

-500 n thick, the intermediate layer with a darker 

ft 

ner side and composed of loosely and longitudinally 



structure 300 



d, slightly 



hyphae 3-3§ n in diameter, many of 












>:-,v 



, • I ft t « t > 

, -v. 






c 



s 




^ 




s' 



Fig. 43. 5. helerosporum . Section X 90; hyaline cystidium, c, colored cystidium, 
c', hyaline spores, s, colored spores s', X 665. 



hich curve into the hymenium and often become there as dark 



d as conduct 



ometimes incrusted: hvme 



mum 



thick, becoming more or less zonate, with cvstidia 



d starting from all parts of the layer, 30-35X6-7 



M 



truding up to 15 u, often colored under 



the deepe 



hymenium; paraphyses filiform, 2 



diameter, branched, numerous at the surface of the hvmenium 



M 



basidi 



9X3^ n, borne 4 to a basidium 



hraceous spores of the same form and dimensions as the 



basidiospores often 
hymenium. 



imbedded throughout the 



1920] 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 221 



Fructifications 5-10 mm. in diameter, becoming confluent 
over areas 1-2 cm. wide and up to 12 cm. long, and reflexed 2-7 
mm. 

On wood and in crevices of the bark of dead limbs and logs of 
Eucalyptus, oak, pecan, and other frondose species. Oregon to 
Mexico. September to April. 

Resupinate specimens of S. heterosporum are not distinguish- 
able in aspect from the darkest colored specimens of S. alboba- 
diurn; all specimens of the former which have been seen so far 
have been bister or seal-brown, which is also the color of the 
upper side of the pileus. Mature specimens of S. hetero- 
sporum differ from those of S. albobadium in the much thicker 
zonate hymenium which has cystidia in all parts of this layer 
and many wholly buried below the surface; the deeper region of 
the hymenium is dark-colored in the type because of the abun- 
dance of dark-colored hyphal ends which are occasionally in- 
crusted, and colored imbedded spores are as numerous as in 
Stereum rugisporum, which has nearly the same geographic 
range. I have not found colored imbedded spores in the collec- 
tion distributed in Ell. & Ev., Fungi Col., 1116, which I refer to 

S. heterosporum on account of other distinctive characters of this 
pecies. 



s 



Specimens examined: 

Exsiccati: Ell. & Ev., Fungi Col., 1116, under the name Stereum 

albobadium. 

Oregon: Portland, C. J. Humphrey, 6125. 

California: Berkeley, C. J. Humphrey, 5981; Campo Mts., 

C. R. Orcutt, 2007, 2008, comm. by U. S. Dept. Agr. Herb.; 
Compton, A. J. McClatchie, in Ell. & Ev., Fungi Col., 1116, 
and (in N. Y. Bot. Gard. Herb., Burt Herb., and Mo. Bot. 
Gard. Herb., 56769); Claremont, D. L. Crawford, 1513, 
comm. by L. O. Overholts, 3325 (in Mo. Bot. Gard. Herb., 
21688) ; Santa Cruz, Dr. Anderson, comm. by W. G. Farlow. 

Arizona: Coronado National Forest, G. G. Hedgcock & W. H. 

Long, comm. by C. G. Humphrey, 2562, 2563 (in Mo. Bot. 
Gard. Herb., 13070, 12811). 

Mexico: Parral, Chihuahua, E. 0. Matthews, 8, and 27, type 

(in Mo. Bot. Gard. Herb., 44282, 44420, 44106); Rosario, 
E. O. Matthews (in Mo. Bot. Gard. Herb.. 44110). 



[Vol. 7 



222 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



73. S. versiforme Berk. & Curtis, Grevillea 1: 164. 



1873; 



Sacc. Syll. Fung. 6: 580. 1888; Massee, Linn. Soc. Bot. Jour. 



27: 193. 1890. 



Plate 6, fig. 73. 



Peniophora Ellisii Massee, Linn. Soc. Bot. Jour. 25: 144. 
1889; Sacc. Syll. Fung. 9: 237. 1891.— An Thelephora obscura 



Persoon, Myc. Eur. 1: 146. 1822 (in 



**** 



Corticium)? 



See 




Peniophora obscura (Pers.) Bresadola, I. R. Accad. Agiati Atti 

111.3:113. 1897. 
Type: in Kew Herb, and Curtis Herb. 

Fructifications at first thin, effused, resupinate, adnate, 
orbicular, becoming confluent, finally thickening, cracking, and 

becoming narrowly reflexed and some- 
what complicate and curling away from 
the substratum, the upper side uneven, 
plicate, somewhat fuscous or blackish; 
hymenium velvety, Prout's brown to 
bister, somewhat papillate; in structure 
200-400 ijl thick, composed of densely ar- 
ranged, ascending and interwoven hyphae, 
some of which are colored; hymenium 
usually simple but sometimes with one or 
two additional zones in some places, con- 
taining heavily incrusted, cylindric cysti- 
dia 45-75X12-24 /*, starting in various 
parts of the hymenium and subhyme- 
nium, wholly buried below the surface of the hymenium or 
emerging up to 15 /x; hy menial surface velvety, with very 
numerous colored paraphyses with bushy-branched tips; spores 

hyaline, even, curved, 5-7X2-3 /x. 

Fructifications 2-10 mm. in diameter, confluent over areas up 
to 7X1-2 cm. ; margin reflexed about 1 mm. usually, rarely up to 

2 mm. 

On the bark of dead limbs of oak, chestnut, birch, and other 
frondose species. Canada to Alabama and westward to Iowa 
and Arkansas. July to February. Common. 

S. versiforme is distinct among the Stereums by its Prout's 
brown, velvety, or at least dull, hymenium, barely reflexed 
margin, and colored, bushy-branched paraphyses, among w T hich 
are scattered large, incrusted cystidia. The presence of these 



Fig. 44. S. versiforme. 
Cystidium, c, and para- 
physes, p, X 665. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 223 



paraphyses, the location of the cystidia in the hymenial side of 
the fructification, and the velvety surface sharply separate 
wholly resupinate specimens of S. versiforme from brownish 
colored forms of Peniophora cinerea. 

Peniophora obscura (Pers.) Bresadola, according to specimen 
collected in Hungary, communicated to me by Bresadola and 
compared by him with an authentic specimen of Persoon, is 
strikingly similar to very young and wholly resupinate specimens 
of Stereum versiforme. There is no European record that P. 
obscura ever has been observed reflexed or has shown any ten- 
dency to become reflexed. In America, S. versiforme is wholly 
resupinate only when very young and soon thickens, becomes 
more or less reflexed, and in well-developed specimens such as 
that cited below, collected by Underwood at White Plains, N. Y., 
has but little in common with P. obscura. For these reasons I 
believe that the name Stereum versiforme should be applied to 
American specimens until Europeans find their Peniophora 
obscura in a reflexed stage identical in its characters with S. 



if 



mens examined 



Exsiccati: Ellis, N. Am. Fungi, 606, under the name Stereum 

papyrinum; Ell. & Ev., N. Am. Fungi, 3209; Ell. & Ev., 
Fungi Col., 611; de Thiimen, Myc. Univ., 307. 

Canada: J. Macoun, 8 in part, 70; on peach tree, J. H. Fault 

(in Mo. Bot. Gard. Herb., 55561). 

Quebec: Hylmer, J. Macoun, 229. 

Ontario: York Mills, J. H. Fault, Univ. Toronto Herb., 322 in 

part (in Mo. Bot. Gard. Herb., 44933). 
New Hampshire: Chocorua, W. G. Farlow (in Mo. Bot. Gard. 

Herb., 55586). 
Vermont: Ripton, E. A. Burt. 
Massachusetts: Arlington Heights, E. A. Burt; Sharon, A. P. 

D. Piquet, comm. by W. G. Farlow (in Mo. Bot. Gard. 
Herb., 55231); Waverly, A. B. Seymour, T 15 (in Mo. 
Bot. Gard. Herb., 18098). 

New York: Alcove, C. L. Shear, 1139, 1304, 1328; East Galway, 

E. A. Burt; Grand View, H. von Schrenk (in Mo. Bot. Gard. 
Herb., 42807); Ithaca, Van Hook, comm. by G. F. Atkin- 
son, 8217; Earner, H. D. House (in N. Y. State Mus. Herb. 



[Vol. 7 
224 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



and Mo. Bot. Gard. Herb., 54354, 54366); White Plains, 
L. M. Underwood (in N. Y. Bot. Gard. Herb, and Mo. 

Bot. Gard. Herb., 5031). 
New Jersey: Newfield, J. B. Ellis, comm. by C. G. Lloyd, and 

in Ellis, N. Am. Fungi, 606, Ell. & Ev., N. Am. Fungi, 3209, 

Fungi Col., 611, and de Thiimen, Myc. Univ., 307. 
Pennsylvania: Michener, type (in Curtis Herb., 4265, and in 

Kew Herb.); Bethlehem, Schweinitz (in Herb. Schweinitz, 

under the name Thelephora amphibola of Schw., Syn. N. 

Am. Fungi, No. 726, but not of Fries); Carbondale, E. A. 

Burt, two collections; State College, C. R. Orton & L. 0. 

Overholts, comm. by L. 0. Overholts, 2661 (in Mo. Bot. 

Gard. Herb., 11419); Trexlertown, W. Herbst, 14- 
Maryland: Glen Sligo, C. L. Shear, 1050, 1095; Hyattsville, 

F. L. Scribner, 90, comm. by U. S. Dept. Agr. Herb.; 

Takoma Park, C. L. Shear, 1020, 1386. 
Virginia: Fairfax, comm. by U. S. Dept. Agr. Herb.; Woodstock, 

C. L. Shear, 1196. 
South Carolina: Salem, Schweinitz (in Herb. Schweinitz, under 

the name Thelephora bufonia of Schw., Syn. N. Am. Fungi, 

No. 725, but probably not T. bufonia Pers., which is too 

imperfectly known for recognition in Europe) ; Summer- 

ville, C. L. Shear, 1227. 
Alabama: Auburn, F. S. Earle (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56785, 56786), and F. S. Earle & 

C. F. Baker (in N. Y. Bot. Gard. Herb, and Mo. Bot. Gard. 

Herb., 56787, 56788). 
Michigan: Ann Arbor, C. H. Kaufman, 21 (in Mo. Bot. Gard. 

Herb., 9808), and Abrams (in N. Y. Bot. Gard. Herb, and 

Mo. Bot. Gard. Herb., 56789). 
Iowa: Woodbine, C. J. Humphrey & C. W. Edgerton, comm 

by C. J. Humphrey, 6518 (in Mo. Bot. Gard. Herb., 20624) 
Missouri: Concordia, C. H. Demetrio (in Mo. Bot. Gard. Herb. 

5030); Oran, H. von Schrenk (in Mo. Bot. Gard. Herb. 

42887); St. Louis, E. A. Burt (in Mo. Bot. Gard. Herb. 

8725) ; Williamsville, B. M. Duggar, 478, 481 . 
Arkansas: Bigflat, W. H. Long, 19783, 19898 (in Mo. Bot. Gard 

Herb., 5921, 9138) ; Cass, W. H. Long, 19800, 19827 (in Mo 

Bot. Gard. Herb., 8636, 8886); Womble, W. H. Long 






19201 



BURT — THELEPHORACEAE OF NORTH AMERICA. XII 



225 



19768, 1987 S, 19881 
5920) . 



Mo. Bot. Gard. Herb., 9143, 8964 



74. S. insigne Bresadola, Nuov. Gior. Bot. Ital. 23: 158* 



1891; Sacc. Syll. Fung. 9: 222. 



1891. 



Plate 6, fig. 74. 



Type: authentic specimen, probably part of the type, in Burt 
Herb. 

Fructification corky, drying rigid, hard, effuso-reflexed, the 
upper surface concentrically sulcate, somewhat zonate, tomen- 
tose, snuff-brown to bister, the recent 
growth at the margin paler; hymenium 
even, pinkish buff to drab-gray and 
pruinose; in structure 1500 m thick, 
with the intermediate layer bordered 
and connected with the tomentum by 
a darker and denser zone and bearing 
on the opposite side a multizonate 
hymenium; hyphae of the intermediate 
layer colored, thick-walled, densely 




Fig. 45. S. insigne. Sec- 
tion of hymenium of authen- 
tic specimen X 665; bottle- 



and longitudinally arranged, 3£ ju in brush paraphyses, p. 
diameter; no cystidia;' paraphyses of 

bottle-brush or aculeate form, numerous and conspicuous in the 
hymenial surface, cylindric, 25-30 X4-4| n] spores published by 
Bresadola as hyaline, even, 4-6X3-3§ /* — none found by me. 

Reflexed l§-4 cm., laterally confluent for 9 cm. in the Florida 
specimen. 

On oak logs. Florida, Venezuela, and Italy. February. Rare. 

This species belongs in the group with S. subpileatum and S. 
septum and is not distinguishable in general aspect from these 
species, but its hymenium contains numerous and conspicuous 
bottle-brush paraphyses and no cystidia, while both of the other 
species named have cystidia. The Venezuelan specimen cited 
below was determined by Berkeley as Stereum illudens, from 
which it appears distinct, for while the type of S. illudens, in 
Kew Herbarium, collected by Drummond, 158, Swan River, 
Australia, has bottle-brush paraphyses for its hymenial surface, 
it has in its subhymenium elongated, cylindric, thick-walled 
organs 6 y. in diameter, up to 100 ju long, a little darker colored 
than the surrounding hyphae and curving outward into the 
deeper portion of the hymenium, which is not zonate. 

10 



226 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



IVou 7 



Specimens examined: 
Italy: Florence, Martelli, comm. by G. Bresadola. 
Florida: C. G. Lloyd, 4846. 
Venezuela: Fendler, 177 (in Curtis Herb.). 



75. S. durum Burt, n. sp 



Plate 6, fie. 75 



Type: in 
Gard. Herb 



Smith, Central Am. Fungi, 147, copy in Mo. Bot 



free 



hard, orbicular, attached by the 



reflexed all around, concentrically sulcate, fuscous to 

bone-brown, with a horn-like crust, 
becoming somewhat shining; hyme- 
nium even, not shining, between pale 




drab 



and tilleul-buff 



somew 




ff 



Fig. 40. S.durum Section reflexed j cm 



pruinose; in structure 2-3 mm. thick, 
hazel throughout, and multizonate or 
stratose, containing many scattered 
crystals, hyphae 3£-4 n in diameter; 
paraphyses of bottle-brush or aculeate 
form, numerous and conspicuous in 
the hymenial surface, cylindric, 12-15 
X4-5 At; no cystidia; no spores found. 
Fructification 3 cm. in diameter. 



of hymenial region of type X 
90; bottle-brush paraphyses, p, 
X 665. 



On dead wood. Mexico. 
S. durum is much thicker, harder 
and more rigid than S. insigne ai 



and 

not tomentose. The microscopic structure agrees exactly with 
that of preparations from an authentic specimen in Kew 
Herbarium of Stereum annosum, No. 99, collected at Neil- 



gherries, Ceylon, and should be 



com 



with the latter 



when better 



For the present the development of a 



3us by S. durum, with characters as stated, is reason for 
;arding this species as distinct from S. annosum. a resuninatft 



ther side of the world 



Specimens 



Exsiccati: Smith, Central Am. Fungi, 147, under the name 

Stereum ferreum. 

Mexico: Jalapa, C. L. Smith, type, in Smith, Central Am. Fungi, 

147. 



1920) 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 227 



76. S.frustulosum(Pers.) Fries, Epicr. 552. 1838; Hym. Eur. 
643. 1874; Morgan, Cincinnati Soc. Nat. Hist. Jour, io: 196. 
1888; Sacc. Syll. Fung. 6: 572. 1888; Massee, Linn. Soc. Bot. 



Jour. 27: 199. 1890. 



Plate 6, fig. 76. 



Thelephora frustulosa Persoon, Syn. Fung. 577. 1801 ; Myc. 
Eur. 1: 134. 1822; Fries, Syst. Myc. 1: 445. 1821.— Thele- 
phora perdix Hartig, Zersetzung. des Holzes, 103-108. pi. 13. 
1878. 

Illustrations: Cooke, Fung. Pests, pi. 20. f. 20; Hartig, loc. 
cit.; Massee, Dis. Cult. Plants, 397. text f. 124; Tubeuf, Dis. 
of Plants, 35. text f.ll, and 430. text f. 260, 261 . 

Fructifications woody, resupinate, tuberculose, crowded as if 
confluent and then broken up into frustules, sometimes grown 
outward from place of attachment and narrowly reflexed or 
with a free margin all around, the upper side black, crust-like, 




1 

\ 



V 





Fig. 47. S . fr ustulosum . Section X 45; bottle-brush paraphyses, p, X 665. 



concentrically sulcate, glabrous; hymenium convex, pinkish 
buff to whitish and pruinose; in structure 800 n or more thick, 
with hyphae densely arranged, radiating outward from the 
place of attachment and bearing a multizonate hymenium in 
which are great numbers of bottle-brush or aculeate paraphyses; 

spores hyaline, even, 5-6X3-3-2- m- 

Fructifications 2-4 mm. in diameter; margin reflexed 3 mm. 
in the best developed specimen known to me. 

On wood of oak logs and stumps in which it causes a pocketed 
or honey-comb rot. Canada to Texas and westward to Oregon, 
in Mexico and in Europe. 

S. frustulosum may be recognized by its occurrence in small 
convex fructifications of woody consistency, crowded together 



» 



[Vol. 7 
228 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



on the under side of dry and hard oak wood or on the sides of 
stumps. On the sides of stumps it may sometimes be found 
reflexed. The bottle-brush paraphyses and many-zoned hy- 
menium are good structural characters for confirmation of the 
determination. 

Specimens examined: 
Exsiccati: Bartholomew, Fungi Col., 1881, 4587; Ellis, N. Am. 

Fungi, 106; Ell. & Ev., Fungi Col., 7; Ravenel, Fungi Car. 

2: 34; de Thiimen, Myc. Univ., 308. 
Sweden : Stockholm, L. Romell, 28; Upsala, E. P. Fries (in Curtis 

Herb.). 
France: Aveyron, A. Galzin, 13935, comm. by H. Bourdot, 

26649. 
Ontario: Carleton Place, J. Macoun, 421 (in Macoun Herb.). 
Vermont: Grand View Mt., E. A. Burt, three collections. 
Massachusetts: Dedham, Hanna; Wellesley, L. W. Riddle, 14- 
New York: Glasco, P. Wilson, 50 (in Mo. Bot. Gard. Herb., 

54763); Ithaca, W. C. Muenscher, 144 (in Mo. Bot. Gard. 

Herb., 56601); Palisades, P. Wilson, 62 (in Mo. Bot. Gard. 

Herb., 54761). 
New Jersey: Alpine, P. Wilson, 8 (in Mo. Bot. Gard. Herb., 

54764); Englewood, P. Wilson, 60 (in Mo. Bot. Gard. 

Herb., 54762); Hackensack Swamp, W. H. Ballou (in 

Mo. Bot. Gard. Herb., 56599); Newfield, J. B. Ellis, in 

Ellis, N. Am. Fungi, 106, in Ell. & Ev., Fungi Col., 7, and de 

Thiimen, Myc. Univ., 308. 
Pennsylvania: Kittanning, D. R. Sumstine. 
Maryland: Hyattsville, F. L. Scribner (in U. S. Dept. Agr. 

Herb.). 
South Carolina: H. W. Ravenel, in Ravenel, Fungi Car. 2: 34; 

Clemson College, P.H. Rolfs, 1621, 1630, 1638. 
Florida: Tallahassee, E. Bartholomew, in Bartholomew, Fungi 

Col., 4587. 
Alabama: Auburn, F. S. Earle & C. F. Baker (in Mo. Bot. Gard. 

Herb., 5079); Montgomery, R. P. Burke, 27 (in Mo. Bot. 

Gard. Herb., 17875). 
Louisiana: A. B. Langlois. 

Texas: Denton, W. H. Long, in Bartholomew, Fungi Col., 1^81; 

Galveston, H. W. Ravenel, 36, comm. by U. S. Dept. Agr. 
Herb. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 229 



Ohio: C. G. Lloyd, 185 (in Lloyd Herb.) ; Loveland, D. L. James 

(in U. S. Dept. Agr. Herb.). 
West Virginia: Paw Paw, C. L. Shear, 1180. 
Kentucky: Crittenden, C. G. Lloyd, 1685. 
Wisconsin: Blue Mounds, Miss A. 0. Stucki, 30; Madison, W. 

Trelease, 83 (in Mo. Bot. Gard. Herb., 44105). 
Iowa: Webster Co., 0. M. Oleson, 450 (in Mo. Bot. Gard. Herb., 

44062) . 

Missouri: Columbia, B. M. Duggar, 1+1+3; Creve Coeur, P 

Spaulding (in Mo. Bot. Gard. Herb., 44103), and E. A 
Burt (in Mo. Bot. Gard. Herb., 7861); St. Louis, Miss C 
Rumbold; Valley Park, E. A. Burt (in Mo. Bot. Gard 
Herb., 44058, 44063). 

Nebraska: Saltillo, C. L. Shear, 1051. 

Kansas: Bourbon Co., A. O. Garrett, 125. 

Oregon: Portland, J. R. Weir, 597 (in Mo. Bot. Gard. Herb. 

36747). 
Mexico: Tepeite Valley, Guernavaca, W. A. & E. L. Murrill 

411 (in Mo. Bot. Gard. Herb., 54545). 
U. S. Northern Pacific Expl. Exp.: Ousmia, C. Wright, comm 

by U. S. Dept. Agr. Herb. 



77. S. roseo-carneum (Schw.) Fries, R. Soc. Sci. Upsal. Actis 
III. 1: 112. 1851. Plate 6, fig. 77. 

Thelephora roseo-carnea Schweinitz, Naturforsch. Ges. Leipzig 
Schrift. 1: 107. 1822 (under C. Corticia). — T. anthochroa Schwei- 
nitz, Am. Phil. Soc. Trans. N. S.4 : 168. 1832, but not T. anthoch- 
roa of European authors. — Corticium lilacino-fuscum Berkeley & 
Curtis, Grevillea 1 : 180. 1873; Sacc. Syll. Fung. 6: 621. 1888; 
Massee, Linn. Soc. Bot. Jour. 27: 143. 1890. — Stereum lilacino- 
fuscum (Berk. & Curtis) Lloyd, Myc. Writ. 5. Letter 68:8. 
1919.— £. sendaiense Lloyd, Myc. Writ. 5. Myc. Notes 48: 680. 
textf. 1015. 1917. — Corticium subrepandum Berkeley & Cooke, 
Grevillea 6: 81. 1878; Sacc. Syll. Fung. 6: 608. 1888; Massee, 
Linn. Soc. Bot. Jour. 27: 119. 1890. 

Illustrations: Lloyd, loc. cit. 

Type: in Herb. Schweinitz, under the name Thelephora 
anthochroa. 

Fructifications coriaceous-soft, thin, usually resupinate, effused, 



[Vol. 7 
230 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



becoming confluent, sometimes with margin barely free, rarely 
distinctly reflexed, with the upper surface tomentose, light buff 
to pinkish buff, the margin entire; hymenium even, cracking in 
a tessellated manner, not shining, light vinaceous purple when 
young, gradually changing to avellaneous when mature; in 

structure 250-300 n thick, composed 
^ . of somewhat longitudinally and 

\Lif *J 'Cp ; •'( [f /") loosely interwoven, hyaline, thin- 

^ S if walled, nodose-septate hyphae 2^-3 







V> 3 (j, in diameter, not differentiated into 

an intermediate layer with a dark 

or dense bordering zone; hymenial 

layer simple when young, with very 

numerous and conspicuous, filiform 
Fig. 48. s. roseo-carneum. Par- paraphyses, colored above and with 

aphysesof type, v; paraphyses, y', ■% , -i * j .• i • i 

of ( ollection at Ithaca, and spores! short - b ™nched tips or bearing short 
s, all x 665. lateral prongs on from 5-20 n of the 

outer portion of the paraphysis, the 
paraphyses less conspicuous when basidia appear; spores white 
in spore collection, even, flattened on one side, 6-9X4-5 m, 
borne 4 to a basidium on simple basidia. 

At first forming little fructifications 3-5 X 2 mm. , which become 
confluent over areas up to 6Xl| cm.; margin becoming free or 
reflexed for 1-3 mm. 

On fallen limbs of frondose species. Canada to North Caro- 
lina and westward to Wisconsin, and in Brazil and Japan. 

Since S. roseo-carneum is nearly always resupinate and does 
not show in sectional preparations of such specimens a distinct 
intermediate layer, its inclusion in the genus Stereum must 
trouble beginners. Fortunately it is a species so unique in 
structure that it may be determined with confidence. Most 
collections are likely to show more or less of the fuscous-lilac 
color, which is intense in young stages; the hymenium cracks 
and has the aspect of Corticium evolvens in other features than 
color, although of different structure; sections of S. roseo- 
carneum show in the hymenial surface filiform paraphyses 
branched above, as shown in the text figure. Such paraphyses 
are present in only one of our Corticiums — Corticium roseum. 
It is regrettable that the Schweinitz type was relabeled by Dr. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 231 



Michener to conform to the name used by Schweinitz in Synopsis 
North American Fungi' and the original label removed from the 
specimen, but Schweinitz gives in the later publication the name 
which he originally used. 

Specimens examined: 
Exsiccati: Ellis, N. Am. Fungi, 515 and 20, the latter under the 

name Corticium incarnatum. 
Ontario: London, J. Dearness, D9Jf5 k, reflexed specimen (in 

Mo. Bot. Gard. Herb., 14251). 
New Hampshire: Chocorua, W. G. Farlow, reflexed specimen; 

North Conway, L. 0. Overholts, 5032, 5161— the latter 

reflexed (in Mo. Bot. Gard. Herb., 56348, 50349). 
Vermont: Middlebury, E. A. Burt, two collections, of which one 

is reflexed; Ripton, E. A. Burt. 
Massachusetts: reflexed specimen, comm. by C. H. Peck; 

Arlington Heights, reflexed specimen, E. A. Burt; Sharon, 

A. P. D. Piguet, comm. by W. G. Farlow. 
Connecticut: C. Wright, type of Corticium lilacino-fuscum (in 

Kew Herb, and Curtis Herb., 5610). 
New York: Alcove, C. L. Shear, 1001, 1002, 1004, 1072, 1321; 

Altamont, reflexed specimen, E. A. Burt; Brookton, W. 

C. Muenscher, 215 (in Mo. Bot. Gard. Herb., 56612) 

Cayuga Lake basin, G. F. Atkinson, 3022; East Galway, 

E. A. Burt; Ithaca, Van Hook, and H. S. Jackson, comm. 
G. F. Atkinson, 8247 and 14396 respectively; North 

Elba, C. H. Kaufman, 13 (in Mo. Bot. Gard. Herb., 16987). 
New Jersey: Newfield, J. B. Ellis, 2487, type of Corticium 

subrepandum (in Kew Herb.), and in Ellis, N. Am. Fungi, 

20, and 515. 

Pennsylvania: Spruce Creek, J. H. Fault, Univ. Toronto Herb., 

312 (in Mo. Bot. Gard. Herb., 44886); State College, L. O. 
Overholts, 2676 (in Mo. Bot. Gard. Herb., 5946), and L. O. 
Overholts & C. R. Orton, comm. by L. O. Overholts, 5041, 
reflexed specimen (in Mo. Bot. Gard. Herb., 56359). 

District of Columbia: Rock Creek, C. L. Shear, 1352; Washing- 
ton, T. Pergande (in U. S. Dept. Agr. Herb.). 

Virginia: Woodstock, C. L. Shear, 786, 788. 

North Carolina: Salem, Schweinitz, type, under the name 

Thelephora anthochroa (in Herb. Schweinitz). 




(Vol. 7 
232 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



West Virginia: Fayette Co., L. W. Nuttall, coram, by Lloyd. 

Herb. 

Michigan: Ann Arbor, C. H. Kauffman, 13. 

Indiana: Crawfordsville, D. Reddick, 9, 10. 

Wisconsin: Palmyra, Miss A. 0. Stucki, 48. 

Brazil: Rio Grande do Sul, Hamburgerberg, G. 0. Malme, 75, 

comm. by L. Romell, 330. 

Japan: A. Yasuda, comm. by C. G. Lloyd (in Mo. Bot. Gard. 

Herb., 55214), and part of type of Stereum sendaiense (in 
Mo. Bot. Gard. Herb., 55448); Sendai, A. Yasuda, reflexed 
specimen (in Mo. Bot. Gard. Herb., 56247). 



SPECIES IMPERFECTLY KNOWN 

Thelephora aculeata Berk. & Curtis, Grevillea i: 149. 1873; 
Sacc. Syll. Fung. 6: 523. 1888. 

The type was collected on the ground in Santee Swamp, South 
Carolina, in June. I had compared with the type a collection 
made by Professor P. H. Rolfs, on the ground, Clemson Col- 
lege, South Carolina, on June 18, and found this collection so 
similar to the type in aspect, although smaller, that I referred 
this specimen to Thelephora aculeata. I had not been able to 
demonstrate basidia for the type nor for the Rolfs specimen; 
now while working out the detailed structure of the latter speci- 
men for publication, I find globose, longitudinally septate 
basidia 9 n in diameter, and hyaline, even spores up to 9X4|-5 
It seems probable that when there is opportunity to examine t 
type again it may be found to have similar basidia and belong 
Tremellodendron. 



/* 



Stereum arenicolum Berkeley in Massee, Linn. Soc 
27: 201. 1890. 



u 



um, effusum, crassum, rieidum. subtus toment 



ferrugineo molli vestitum; hymenio levi, glabro fusco-purpu- 
rascente; sporae ellipsoideae, 7X4-5 n (Berk, in Herb. n. 3822). 

"On sand under trees, Vera Cruz. 

"Rigid, thick, 2-3 inches across, attached to the sand and 
probably decayed wood by a dense ferrugineous tomentum; 
margin sometimes slightly upraised; substance pale cinnamon." 

The above should be compared with S. crassum. 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 233 



\ 



Stereum cuneatum Lloyd, Myc. Writ. 4. Letter 54: 7. 1916. 

"Pileus cuneate, tapering to the base (2 cm. high), cut into 
a few fimbriate segments. Surface pale, smooth. Hymenium 
unilateral, pale yellow (honey yellow of Ridgway), smooth. 
Cystidia none. Spores globose, 3§-4 mic, hyaline, smooth. 
The plant grows densely caespitose in the earth, from a common 
mycelial base. It belongs in Section 7 of my recent pamphlet 
on Stipitate Stereums." Florida. 

Perhaps the above is S. Burtianum or S. tenerrimum. 



Stereum cupulatum Patouillard in Duss, Fl. Crypt. Antilles 
Fr. 233. 1904. 

Scattered or close together, orbicular, from resupinate 
becoming cup-shaped, attached by a dorsal point, coriaceous, 
rigid, hard; external face glabrous, not zonate, brown, the 
margin entire or sinuate, acute; hymenium pruinose, even, 
concave, dull cinereous, reddish towards the border; trama 
compact, brown-umber; spores cylindric-ovoid, colorless, 6X3 ft; 
no cystidia. 

Fructifications 6-8 mm. in diameter. 

On bark of Prunus Dussii. — Forest of Buins-Jaunes. Duss, 
212. 

The above is a translation of the original description; the 
species seems to be very near, if at all distinct from, Stereum 
vibrans, which Patouillard did not recognize among the species 
of Guadeloupe. 



Stereum fragile Patouillard, Soc. Myc. Fr. Bui. 16: 179. 
1900; Sacc. Syll. Fung. 16: 187. 1902. 

Fructification resupinate at first, becoming dimidiate, orbic- 
ular, rigid, hard, more or less incised at first, the margin erect 
and acute; upper surface plane, ochraceous russet, tomentose, 
with some reddish and nearly glabrous concentric zones; trama 
1 mm. thick, whitish, compact; hymenium plane, livid, becom- 
ing purplish; cystidia abundant, fusoid, not colored, thin-walled, 

40X10/X- 

On decaying wood. Guadeloupe. 

This fungus is very fragile and divides radially with great 

ease. Its aspect is like that of S.fasciatum, S. lobatum, etc., but 



[Vol. 7 
234 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



it is easily distinguished by the violaceous tint of the hymenium. 
I have not seen authentic specimens of S. fragile, but from the 
foregoing translation of the original description, it seems very 
probable that S. fragile may prove a synonym of S. albobadium, 
a species common in the West Indies but not recognized by 
Patouillard among the species of Guadeloupe. 



Stereum fimbriatum Ellis, Torr. Bot. Club Bui. 6: 133. 1877. 

According to the authentic specimen from Ellis to Cooke in 

Kew Herb., this is a whitish, flaxy mass having no hymenium 

and quite indeterminable. 



Stereum Galeottii Berkeley, Hooker's Jour. Bot. 3: 15. 1851; 
Sacc. SylL Fung. 6: 574. 1888; Massee, Linn. Soc. Bot. Jour. 
27: 176. 1890. 

"Umbonato-sessile, parvum, convexum, rigidum; pileo cer- 
vino velutino-tomentoso crebrissime badio-zonati; zonis hie illic 
glabris nitentibus; hymenio cinereo-alutaceo. Galeotti, No. 
6853. 

"Hab. Caripi, Spruce; Vera Cruz, Galeotti; Xalapa, Mr. 
Harries. 

"Pileus l\ inch broad, 1 inch long, subflabelliform, umbonato- 
sessile, mostly convex above, slightly undulated, thin but rigid, 
fawn-colored, clothed with velvety down; repeatedly zoned; 
zones mostly very close and narrow, frequently forming bay- 
brown, smooth and shining, alternating with paler fasciae. 
Hymenium tan-colored with a cinereous tinge. 

"Undoubtedly nearly allied to Stereum lobatum, Kze, but a 

much smaller and neater species/' 

The type of the above should be compared with Stereum 
versicolor. 



Stereum griseum Schweinitz, Naturforsch. Ges. Leipzig 
Schrift. 1: 106. 1822 (under B. Sterea of Thelephora); Fries, 
Elenchus Fung. 1: 179. 1828. — Stereum porrectum Fries, Epicr. 
548. 1838; Sacc. SylL Fung. 6: 579. 1888. 

I have been unable to find any Schweinitzian specimen of this 
species. It seems probable that the description was based on 
the old stage of Stereum fasciatum in which the attachment is by 






1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 235 



umbo prolonged into stem-like form. Such fructifications occur 
rarely and are perplexing if not gathered in the same collection 
with the usual sessile fructifications. 



S. ochroleucum Fries, Hym. Eur. 639. 1874; Sacc. SylL 
Fung. 6: 562. 1888; Massee, Linn. Soc. Bot. Jour. 27: 184. 

1890. 

Corticium ochroleucum Fries, Epicr. 557. 1838. — Not Stereum 

ochroleucum Bres. Ann. Myc. 1: 91. 1903, nor Brinkmann, 
Westfalische Pilze, 49. 

Type: authentic specimen in Kew Herb. 
This species does not occur in North America and adjacent 
regions although reported from time to time from United States, 
Cuba, and Venezuela. Since I have not received under any 
name specimens of the true Stereum ochroleucum from European 
correspondents, this species is probably rare in Europe, and it 
may help toward recognition of the species to call attention to 
the specimen in Kew Herbarium. 
The specimen is labelled: 

"Corticium ochroleucum Fr. 
Svex. Westm. 
Maji — leg. Lbd." 

This specimen agrees well with the original description; its 
reflexed portion is \\ cm. broad, about 1-1 1/5 mm. thick as the 
sections show in my preparation; the consistency is soft in com- 
parison with S. hirsutum and the hyphae about 2\ mm. in 

diameter, granule-incrusted, and interwoven throughout the 
thickness of the pileus rather than parallel and longitudinally 
arranged side by side as in S. hirsutum and S. sulphur atum . In 
other words there is not the sharply marked intermediate layer 
which Fries regarded as an important distinctive character of 
the genus Stereum, and this is probably the reason for his orig- 
inally regarding this species as a Corticium although broadly 
reflexed. There is not present a hardened crust or golden zone 
to mark the upper side of the intermediate region, but instead 
the hyphae become more loosely arranged toward the surface and 
become the hairy covering of that side. No cystidia, gloeo- 
cystidia, nor colored conducting organs are present; the spores 
are hyaline, even, 4^-5 X 3 p. 



[Vol. 7 
236 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



The American Stereum spumeum has aspect and structure very 
similar to Stereum ochroleucum Fr. but differs by having incrusted 
cystidia. 



Stereum unicum Lloyd, Myc. Writ. 4. Stip. Stereums, 35. 

text f. 556. 1913. 

The type is in New York State Museum under the name 
Thelephora speciosa unless relabeled to conform to the name 
applied by Lloyd. The type bears no basidia yet and is not 
determinable as to genus; it was collected in Providence, 
Saratoga County, New York, where I have been looking for a 
fertile specimen when in the original locality occasionally in the 
summer. 



EXCLUDED SPECIES 



Stereum acerinum (Pers.) Fr. is Aleurodiscus acerinus (Pers.) 
v. Hohn. & Litsch. 

Stereum acerinum var. nivosum Berk. & Curtis is Aleuro- 
discus nivosus (B. & C.) v. Hohn. & Litsch. 

Stereum calyculus Berk. & Curtis is Craterellus calyculus 
(B. & C.) Burt. 

Stereum candidum Schweinitz is Aleurodiscus candidus 
(Schw.) Burt. 

Stereum carolinense Cooke & Ravenel is Sparassis spathu- 

latus (Schw.) Fr. 

Stereum duriusculum, as determined by Patouillard in Duss, 
Fl. Antilles Fr. 232. 1903, is probably Hypochnus pallescens 
(Schw.) Burt, a species common in the West Indies. 

Stereum Guadelupense Patouillard, Soc. Myc. Fr. Bui. 15: 

201. pi 10. f. 1. 1899. According to von Hohnel & Lit- 
schauer, K. Akad. Wiss. Wien Sitzungsber. 116: 753. 1907, 
this is a Boletus overrun by a Sepedonium. 

Stereum Haydeni Berkeley in Massee, Linn. Soc. Bot. Jour. 
27: 199. 1890. 

The type, in Kew Herbarium, was collected in Ohio; it is 
strictly resupinate, has its hyphae loosely interwoven from 
hymenium to substratum, and has no characters which justify 
its inclusion in Stereum as comprehended in my work. The 



1920] 

BURT — THELEPHORACEAE OF NORTH AMERICA. XII 237 



hymenium is deteriorated but shows no cystidia; the species 
may be sought for in Ohio as a probable Corticium. 

Stereum insolitum Lloyd, Myc. Writ. 5. Myc. Notes 47: 665. 
text f. 956. 1917, is a young specimen of Thelephora regularis 
Schw. 

Through the kindness of Professor McFarland, I have exam- 
ined his portion of the original specimen. Most of the spores 
attached to the basidia are as published by Lloyd; a few spores 
are 6-7X5 /*, rough-walled and still hyaline; occasional spores in 
a preparation from near the base of the pileus are colored and 
tuberculate-irregular. 

Stereum Leveillianum Berk. & Curtis is Tremellodendron 
Leveillianum (B. & C. )Burt. 

Stereum Micheneri Berk. & Curtis is Thelephora albido- 

brunnea Schw. 

Stereum Mancianus Sacc. & Cub. is Aleurodiscus strumosus 
(Fr.) Burt. 

Stereum populneum Peck, N. Y. State Mus. Rept. 47: 145. 
1894. 

This is known in resupinate form only and should not be 
included in Stereum. 

Stereum pruinatum Berk. & Curtis, Linn. Soc. Bot. Jour. 10: 
332. 1868. 

This is known in resupinate form only and should not be 
included in Stereum. 

Stereum scriblitum Berk, & Cooke, Grevillea 7: 102. 1879; 
Sacc. Syll. Fung. 6: 567. 1888. 

The type collected by Gerard, 171 (in Kew Herb.) was studied. 
This is the conidial stroma of Ustilina vulgaris. 

Stereum seriatum Berk. & Curtis is Aleurodiscus seriatus 
(B. & C.) Burt. 

Stereum spongiosum Massee is Thelephora albido-brunnea 
Schw. 

Stereum strumosum Fries is Aleurodiscus strumosus (Fr.)Burt. 

Stereum subcruentatum Berk. & Curtis, Am. Acad. Arts & 
Sci. Proc. 4: 123. 1858, is Aleurodiscus subcruentatus (Berk. 
& Curtis) Burt, n. comb.; now included among American spe- 
cies, because of collections received from California and Oregon. 



238 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Stereum triste Berk. & Curtis, Linn. Soc. Bot. Jour. 10: 332 



1868. 



This is the conidial stroma of a Pyrenomycete and shows 
young perithecia under the stroma in the type in Curtis Herb. 
Collection in Kew Herb., C. Wright, 252, has similar structure 
but did not show perithecia in my sections. 

(To be continued.) 



[Vol. 7, 19201 



240 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Explanation of Plate 

PLATE 2 

All figures of plates 2-6 have been reproduced natural size from photographs of 
dried herbarium specimens unless otherwise noted. 

Fig. 1. Stereum caperatum. Specimen collected at St. Martin ville, La., by A. B. 

Langlois. 

Fig. 2. S. hydrophorum. Specimen collected at Rio Mato, Venezuela, by M. A. 

Carriker. 

Fig. 3. S. Ravenelii. Type distribution in Ravenel, Fungi Car. 4: 13. 

Fig. 4. S. surinamense. Specimen collected at Consuelo, San Domingo, by N. 

Taylor, 12. 

Fig. 5. S. Burtianum. Specimens collected at Amherst, Mass., by P. J. Anderson. 

Fig. 6. S. quisquiliare. From Lloyd's illustration of the type. 

Fig. 7. S. aurantiacum. Specimens collected at Port Antonio, Jamaica, by F. 

S. Earle. 

Figs. 8 and 9. S. diaphanum. Fig. 8 from type of S. diaphanum, and Fig. 9 

from type of S. Willeyi. 

Fig. 10. S. exiguum. Type. 
Fig. 11. S. tenerrimum. Type. 



pergamenum 



Type distribution in Ravenel, Fungi Car. 3 : 25. 






Ann. Mo. Bot. Gard., Vol. 7, 1920 



Plate 2 




BURT— THELEPHORACEAE OF NORTH AMERICA 



1 8TEREUM CAPERATUM— 2. 8. HYDROPHOHUM.— 3. S. RAVENELII— 4. S. SURINAMENSE. 
5. 8. RURTIANUM— <>. S. QUISQUILIARE— 7. S. AURANTIACUM.— 8-9. S. DIAPHANUM.— 10. S. EXI 

GUUM.— 11. S. TENERRIMUM.— 12. S. P :RGAMENUM. 



Vol. 7, 1920] 
242 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Explanation of Plate 

PLATE 3 

Figs. 13 and 14. S. pallidum. Fig. 13, specimen collected and determined by G. 
Bresadola; Fig. 14, specimen collected at Blowing Rock, N. C, by G. F. Atkinson. 

Fig. 15. S. elegans. Specimen collected at Mayaguez, Porto Rico, by B. L. 
Santiago, 12. 

Fig. 234. S. decolorans. Type. 

Fig. 16. S. radicans. Specimen collected at Grenada, by W. E. Broadway. 
Fig. 17. S. pusiolum. Specimen collected at Rio Piedras, Porto Rico, by J. R. 
Johnston, 89. 

Fig. 18. S. glabrescens. Specimen collected at Sumidero, Cuba, by J. A. Shafer, 
13906. 

Fig. 19. S. fissum. Type. 

Fig. 20. S. cTjphelloides . Type. 

Fig. 21. S. Hartmanni. Specimen collected at St. Kitt's, by N. L. Britton & 
J. F. Cowell. 

Fig. 22. S. craspedium. Specimen collected in Dutch Guiana, by J. Samuels. 
Fig. 23. S. petalodes. From C. G. Lloyd's illustration of the type. 



Ann. Mo. Bot. Gard., Vol. 7, 1920 



Plate 3 




BURT— THELEPHORACEAE OF NORTH AMERICA 

13-14. STEREUM PALLIDUM— 15. S. ELEGAN8— 234. S. DECOLORANS.— 10. S. RADIOANS — 17 
S. PUSIOLUM.— 18. S. GLABRKSCENS— ID'S. FISSUM.— 20. S. CYPHELLOIDES.— 21. S. HARTMANNI.- 

22. S. CRASPEDIUM.— 23. S. PKIALODKS 



244 



[Vol. 7,1920] 
BOTANICAL GARDEN 



Explanation of Plate 



PLATE 4 



Fig. 24. S. proliferum. Type. 
Fig. 25. S. caespilosum. Type. 
Fig. 26. S. fuscum. Specimen c 



Middlebury 



Fig. 27. S. rufum. Specimen collected at Middlebury, Vt., by E. A. Burt. 

Fig. 28. S. Pini. Specimen collected at Chocorua, N. H., by W. G. Farlow, 37. 

Fig. 29. S. purpureum. Specimen collected at North Ferrisburg, Vt., by E. 
Burt. 

Fig. 30. S. rugosiusculum. Specimen collected at Creve Coeur Lake, Mo., 
E. A. Burt. 

Figs. 31 and 32. S. Murrayi. Fig 31, old reflexed specimen collected at Gra 
View Mt., Vt., and Fig. 32, resupinate specimen collected at Ripton, Vt., both by 
A. Burt. 

Fig. 33. S. saxitas. Type. 

Figs. 34 and 35. S. styracifluum. Fig. 34, type; Fig. 35, specimen collected 
Auburn, Ala., by F. S. Earle & C. F. Baker. 

Fig. 36. S. gausapatum. Specimen collected at Toronto, Canada, by T. Langt 

Fig. 37. S. australe. Type. 

Figs. 38 and 39. S. rugosum. Fig. 38, specimen collected at Ithaca, N. Y. 



E. A. Burt. 



specimen collected 



Ann. Mo. Bot. Gard., Vol. 7, 1920 



Plate 4 




HURT— THELEPHORACEAE OF NORTH AMERICA 



24. STKKKTM PROUFEKtM — 2f> S CAESPITOBUM,— 26. S. FUSCUM.— 27. S. RUFUM.— 28. S. PINI. 
29. S. PURPUREUM.— 30. 8. RUGOSIUSCULUM.— 31-32, S. MURRAY!.— 33, a SAXITAS.— 34-35. S. STY 

RACIFLUUM,— 36, S. GAUSAPATUM.— 37 S. AUSTRALE.— 38-39. S. RUGOSUM. 



■ 



[Vol. 7, 1920 J 
246 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Explanation of Plate 

plate 5 

Fig. 40. S. sanguinolentum. Specimen collected in Little Notch, Vt., by E. A. 



Burt. 



Fig. 41. S. sulphuratum. Specimen collected at Auburn, Ala., comm. by F. S. 

Earle. 

Fig. 42. S. hirsutwn. Specimen collected at Smugglers Notch, Vt., by E. A. Burt. 

Figs. 43-45. S. fasciatum. Fig. 43, young effuso-reflexed stage, and Fig. 44, old 
stage with attachment by umbos, both collected at Middlebury, Vt., by E. A. Burt; 
Fig. 45, specimen collected at Formosa, Japan, by S. Kusano, II. 16. 

Fig. 46. S. lobatum. Specimen collected at Lake City, Fla., by P. L. Ricker, 893. 

Fig. 47. S. versicolor. From Berkeley's illustration of the type. 

Fig. 48. S. rameale. Specimen collected at Arlington, Mass., by E. A. Burt. 

Fig. 49. S. sericeum. Specimen collected at Middlebury, Vt., by E. A. Burt. 

Fig. 50. S. pubescens. Type. 

Fig. 51. S. conicum. Type. 

Fig. 52. S. vibrans. Specimen collected at Rose Hill, Jamaica, by F. S. Earle, 303. 

Fig. 53. S. radiatum. Specimen collected at Harraby, Ontario, by E. T. &. S. A. 

Harper, 636. 
Fig. 54. S. patelliforme. Type. 
Fig. 55. S. ochraceo-flavum. Specimen collected at Albany, N. Y., by H. D. 

House. 
Fig. 56. S. abietinum. Specimen collected at Smugglers Notch, Vt., by E. A. 

Burt. 
Fig. 57. S. ambiguum. Specimen collected at Ripton, Vt., by E. A. Burt. 



Ann. Mo, Bot. Card., Vol. 7, 1920 



Plate 5 




BURT— THELKPHORACEAE OF NORTH AMERICA 

40. 8TEREUM SANGUINOLENTUM.— 41. 8. SULPHURATUM.— 2. S. HIRSUTUM.— 43-45. 8. FASCIA 
TUM— 46. S. LOBATUM.— 47. 8. VERSICOLOR.— 48. S. RA ME ALE. —49. S. SERICEUM.— 60. S. PUBE 
8CENS.— 51. 8. CONICUM.— 52. S. VIBRANS — 53. 8. RADIATUM.— 54. S. PATELLIFORME.- 

OCHRACEO-FLAVUM.— 56. S. ABIETINUM— 57. S. AMBIGUUM. 



55. S 



[Vol. 7, 1920] 
248 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Explanation of Plate 

PLATE 6 
Fig. 58. S. rugisporum. Specimen collected at Flagstaff, Ariz., by W. H. Long, 



21307. 

Fig. 59. S. umbrinum. Specimen reflexed on both sides, collected at Valley Park, 
Mo., by E. A. Burt. 

Fig. 60. S. papyrinum. Specimen on under side of a small limb and reflexed on 
both sides, collected at Alto Cedro, Cuba, by Underwood & Earle, 1481. 
Fig. 61. S. Earlei. Type. 

Fig. 62. S. Chailletii. Reflexed specimen collected at Albuquerque, N. M., by 
. H. Long & P. W. Seay, 21313. 

Fig. 63. S. ferreum. Reflexed specimen collected at Cinchona, Jamaica, by W. 



W 



A. & E. L. Murrill, 458. 

Fig. 64. S. cinerascens. Specimens collected at Middlebury, Vt., by E. A. Burt. 
Fig. 65. S. magnisporum. Type. 

Fig. 66. S. spumeum. Specimen collected at Cordoba, Mexico, by W. A. & E. L. 
Murrill, 1214. 

Fig. 67. S. erumpens. Type. 

Fig. 68. S. sulcatum. Type. 

Fig. 69. S. subjrileatum. Specimen collected at St. Martinville, La., by A. B. 
Langlois. 

Fig. 70. S. se-pium. Type. 

Fig. 71. S. albobadium. Specimen collected at Seven Locks, Md., by P. L. Ricker, 
1007. 

Fig. 72. S. hetcrosporum. Type. 

Fig. 73. S. versiforwe. Specimen collected at White Plains, N. Y. by L. M. 
Underwood. 

Fig. 74. S. insigne. Specimen collected in Florida by C. G. Lloyd, 4846. 

Fig. 75. S. durum. Type. 

Fig. 76. S. frustulosum. Specimens collected at Creve Coeur, Mo., by E. A. Burt. 

Fig. 77. S. roseo-carneum. Specimen collected at Arlington Heights, Mass., by 
E. A. Burt. 



Ann. Mo. Bot. Card., Vol. 7, 1920 



Plate P> 




BURT— THELEPHORACEAE OF NORTH AMERICA 



58. STEREUM RUGISPORUM.— 59. 8. UMBRINUM.— 60. 8. PAI'YRINUM.— 61. S. EARLEL— 62. S. 
CHAILLETIL— 63. S. FERREUM.— 64. S. CINERASCENS — (if,, s. M VGNISPORUM.— 66. S. SPUMEUM.- 
67. S. ERUMPENS.— 68. S. SULCATUM.— 69. S. SUBPILEATUM.— 70 S. BEPIUM.— 71. S. ALBOBADIUM. 
72. S. HETEROSPORUM.— 73. S. VERSIFORME.— 74. S. INSIOXI .— 75. S. DURUM.— 76. S. FRUSTU- 

LO.sr.M.— 77. 8. ROSEO-CARNKIM. 



Annals 

of the 

Missouri Botanical 



Garden 




Vol. 7 



NOVEMBER, 1920 



No. 4 



STUDIES IN THE PHYSIOLOGY OF THE FUNGI 
XL Bacterial Inhibition by Metabolic Products 

WILLIAM H. CHAMBERS * 

Formerly Rufus J. Lackland Fellow in the Henry Shaw School of Botany of 

Washington University 

Considerable work has been done on the early phases of 
growth of bacteria in liquid media. Rahn ('06), Coplans ('07), 
Penfold ('14), Chesney ('16), Salter ('19), and others have 
shown quite definitely the factors involved in the lag phase 
of growth preceding the phase of logarithmic increase. They 
have demonstrated that the lag can be eliminated if the trans- 
fers are made during the period of logarithmic increase, but 
that certain factors such as difference in temperature, composi- 
tion of the medium, or the age of the culture will produce a 
latent period immediately following the transfer. 

Data on the later growth periods of bacteria are less extensive. 
Based on the total number of viable bacteria, in the culture, the 

growth curve can be traced roughly as follows: It rises abruptly 
at first, which is the phase of logarithmic increase, then ascends 
more gradually until the peak is reached, and finally descends 
until the culture is sterile. The influence of inhibitory factors 
is most clearly seen in the later periods, those following the phase 
of logarithmic increase, the study of which is of fundamental 
and practical importance both in killing pathogenic bacteria, 
that is, hastening the decline in the growth curve, and in pro- 
longing the life of useful cultures, suspending this decline. In 
the work presented here, emphasis is placed on the later periods 
of growth and on the influence of the products of a growing cul- 
ture on the path of the growth curve. 



Ann. Mo. Bot. Gard., Vol. 7, 1920 



(249) 



IVol. 7 



250 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Literature 

The literature on the subject of the inhibition of bacteria in 
culture by their own metabolic products is widely scattered, 
and the investigational work in this phase of growth studies 
is very meager. The entire subject is often dismissed with 
some such statement as "the organisms are finally killed by 
their own products." 

From time to time different investigators have sought to 
determine if there is a special metabolic product, enzymatic in 
nature, which inhibits the growth of the organism producing 

it. One of the earliest publications on this subject appeared by 
Eijkmann ('04). He grew Bacillus coli in gelatin at 37° C, 
treated it in different ways, and then solidified and reinoculated 
the gelatin. He concluded that Bacillus coli in gelatin pro- 
duced a diffusible, thermolabile substance which would not pass 
through a porcelain filter and which inhibited growth of Bacillus 
coli and other organisms, for treatment with ether, subjection 
to heat, or filtration through a porcelain filter removed some 
inhibiting substance and permitted a streak growth on the 
solidified gelatin. 

The following year Conradi and Kurpjuweit ('05,'05 a ) extended 
the work of Eijkmann, finding the same action in bouillon. 
They called the substance "autotoxin" and applied the theory 
to the germicidal action found in feces. They reported that 
the "autotoxin" of Bacillus coli was killed by boiling but was 
virulent up to a dilution of 1:3200 in a 10-hour culture, and 
that the heated stool filtrate from a paratyphoid patient would 
support growth of the same Bacillus paratyphosis in a 1 : 50 dilu- 
tion, but the unheated filtrate only in a 1 : 400 dilution. Roily 
('06), Passini ('06), and Manteufel ('07) disputed the findings 
of Eijkmann, also those of Conradi and Kurpjuweit, and held 
that the existence of inhibitory substances had not yet been 
proved. Roily could not repeat the work of Conradi and Kurp- 
juweit with the same results but found that the filtered half 
of a 20-hour bouillon culture gave better growth than the cooked 
half. Manteufel claimed that the loss of necessary food ma- 
terial from the media explained some of the results attributed 
to "autotoxin." Kruse ('10) summarized these reports and 



I 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 251 



explained the death of organisms in culture as probably due 
rather to the exhaustion of the media and the accumulation of 
well-known metabolic products than to an "autotoxin." He 
suggested the possibility of the exhaustion of the media and 
the accumulation of products causing the dea:h of a few of the 
weaker individuals, which become self-digested, thereby releas- 
ing previously formed "autotoxin." Acids and alkalis are re- 
ported by him as inhibitory agents, although bouillon in which 
pneumococci had grown would not support a second growth 
even on readjusting the reaction. 

In connection with some work on the latent period of growth, 
Chesney ('16) found that pneumococci in plain broth showed 
marked inhibition 24 hours after inoculation, the number of 
bacteria decreasing rapidly to zero, but if after 96 hours a por- 
tion of the bouillon was filtered through a porcelain filter and 
reinoculated, no inhibition was evident, indicating that the 
inhibitory substance was kilted or attenuated i:i 3 days at 37° C. 

It is apparent from the literature cited above that results 
are conflicting concerning the production of an enzymatic "auto- 
toxin," and while the reports favoring the existence of such 
a product are not conclusive, no other satisfactory explanation 
for the observed reactions has been demonstrated. 

Recent literature has indirectly contributed considerable of 
value concerning the relationship of acid and alkali to growth 
and death of bacteria, through the more general use, since 1916, 
of the hydrogen ion concentration as an expression of acidity 
of media. Winslow and Lochridge ('06), working on Bacillus 
coli and Bacillus typhosus, stated that the toxic effect of inor- 
ganic acids, HC1 and H 2 SO<, corresponded to their dissociation, 
but with organic acids, acetic and benzoic, the undissociated 
molecule was also important, for results did not correspond to 
the dissociation of the acids. Michaelis ('14) advanced the 
idea that organisms produce acid to a certain concentration, 
which he found to be Ph 5.0 with Bacillus coli in lactose bouillon, 
and that they automatically protect themselvas against harm- 
ful amounts. 

Since that time a great deal has been published on final or 
limiting hydrogen ion concentrations for different organisms, 



(Vol. 7 



252 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



but only a very little on the effect on growth of changes in hy- 
drogen ion concentration during growth. Clark ('15) deter- 
mined the final P H of 16 cultures of Bacillus coli in 1 per cent 
dextrose medium as P H 4.67-5.16, and Clark and Lubs ('15) 
in constructing their media for differentiating the members of 
the colon-aerogenes group showed that a reversion of reaction 

toward the alkaline may take place, depending on the dextrose, 
but they did not show the relationship between reversion of 
reaction and growth. Itano ('16) reported that with Bacillus 
sublilis, Streptococcus erysipelatus, and Streptococcus lacticus in 
plain broth, acid was formed in alkaline media and alkali in 
the acid media, thus bringing the P H to a certain definite hy- 
drogen ion concentration. Fred and Loomis ('17) showed a 
wide range of reaction for Bacillus radicicola, obtaining good 
growth between P H 3.9 and 11.1. They also demonstrated that 
the hydrogen ion concentration approaches the neutral point 
during growth. Shohl and Janney ('17) found that P H 4.6- 
5.0 was inhibitory for Bacillus coli in urine. Ayers, Johnson, 
and Davis ('18) added streptococci to the list of organisms whose 
final P H was demonstrated. They separated the pathogenic 
from the non-pathogenic forms on the basis of limiting hydrogen 
ion concentration, the former reaching P H 5.4-6.0 and the latter 
P„ 4.6-4.7. 

The work of Ayers and Rupp ('18) on simultaneous acid and 
alkali fermentations showed some interesting P H curves. They 
found in a .5 per cent dextrose medium that Bacillus coli pro- 
duced acid to P H 4.8 but that Bacillus aerogenes produced less 
initial acid and the reaction reverted to P H 6.5. From quan- 
titative determinations of dextrose and of formic, acetic, lactic, 
and succinic acids, they explained the reversion of Bacillus 
aerogenes as a fermentation of the organic acids, mostly formic 
and acetic, to carbonates. With the alkali-forming milk bac- 
teria, they showed alkaline fermentation of citrate, acid fer- 
mentation of dextrose, and a practically neutral reaction from 
the simultaneous fermentation of the citrate and dextrose. 
Gillespie ('18) found Actinomyces chromogenus gave a poor 
growth at P H 4.8-5.2 and decreased the hydrogen ion concen- 
tration of the media during growth. Wyeth ('18) showed with 
Bacillus coli in glucose bouillon that the final P H varied with 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 253 



the initial P H ; i. e., P H 7.11 progressed to P H 5.70 and P H 4.96 
went to P H 4.68. He also showed a difference in critical P H 
according to the acid used, whether hydrochloric, acetic, or 
lactic acid. Wyeth ('19) extended his previous work and found 
in 2 per cent peptone that an initial range of P H 4.29-9.37 
gave a final range after 216 hours of P H 5.92-8.55, and that 
with an initial P H above 8.48 the production of acid exceeded 
that of alkali and the reaction approached P H 8.48. Indole 
formation was completely inhibited by dextrose, partially by 
sucrose, and not at all by starch. Avery and. Cullen ('19) used 
the final hydrogen ion concentration to separate strains of 
Streptococcus hemolyticus; 124 human strains attained a final 
P H of 4.8-5.3, while 40 dairy strains reached P H 4.3-4.5. 

Considerable work has recently appeared on the pneumo- 
coccus. Cullen and Chesney ('18) showed the relation of the 
growth of pneumococcus in plain broth to hydrogen ion con- 
centration. The bacteria increased to 420,C 00,000 per cc. in 
13.8 hours and then decreased to 160 per cc. in 96 hours. The 
hydrogen ion concentration increased from F v 7.70 to P H 7.03, 
but these investigators expressed the opinion that the increase 
in hydrogen ion concentration is not the sole cause of the cessa- 
tion of growth. Avery and Cullen ('19 a ) showed some inter- 
esting reactions of pneumococcus to carbohydrates. One per 
cent of maltose, saccharose, lactose, galactose, raffinose, dex- 
trose, or inulin produced a final P H of about 5.0. With .4 per 
cent dextrose, as high an hydrogen ion concentration was at- 
tained in 48 hours as with 1 or 2 per cent dextrose. Pneu- 
mococcus differed from Bacillus coli in that it produced acid in 
plain broth, and growth ceased at about P H 7.0. When this 
culture was readjusted to P H 7.8 and reinocalated, no growth 
occurred unless carbohydrate was added, yet the filtrate from 
a dextrose culture at P H 5.2 if readjusted to P H 5.8, 7.0, or 8.0 



would 



Growth could onlv be initiated 



within certain limits, in carbohydrate media P H 8.3-6.8 
and in plain broth P H 8.1-7.0. They concluded that the 
exhaustion of fermentable carbohydrate is only one of the many 

1 To avoid confusion, attention is called to the distinction between the concen- 
tration of the hydrogen ions and the symbolic P H . A numerical increase in hy- 
drogen ions is expressed as a decrease in terms of Ph. 



254 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



(Vol. 7 



factors involved in the complex phenomenon of growth inhi- 
bition. 

Lord and Nye ('19) have demonstrated the relation of time 
to inhibitory action of hydrogen ion concentration with pneu- 
mococcus. They found that Pneumococcus Type I was killed 
in 1 hour at P H 4.5-4.7, in 3 hours at P H 5.3, and in 6 hours 
at P H 6.15, but survived 6 hours at P H 6.35, and that between 
P H 6.8 and P H 5.1 there was a direct relation between the P H 
and the time required for the death of the pneumococcus. In 
mixtures of equal quantities of emulsions of washed pneumo- 
cocci and buffer solutions of different hydrogen ion concentra- 
tions they observed very little dissolution of the bacterial cells 
between P H 8.0 and P H 7.0 or between P H 5.0 and P H 4.0, but 
noticed almost complete dissolution in the zone of P H 6.5- 5.5. 

Bunker ('19) published the results of investigations of Bacillus 
diphtheriae extending over several years. The hydrogen ion 
curves in sugar-free and in 1 per cent dextrose media agree 
very closely with those of Bacillus coli in the experimental work 
of this report. He also showed that toxin was only produced 
within a rather narrow hydrogen ion range, P H 7.8-8.25. 
The best growth, measured by pellicle formation, was obtained 
when the initial reaction was P H 7.3-7.5. Cohen and Clark 
('19) investigated the effect of hydrogen ion concentration on 
the rate of growth of different organisms during the early part 
of the growth curve, the period of logarithmic increase. Cul- 
tures were inoculated into media adjusted over a wide range of 
varying initial hydrogen ion concentrations, and observed for 
the first 10 hours of growth. In general, the different organ- 
isms reacted similarly. The most marked effect of the hydrogen 
ion concentration on early growth was found near the critical 
acid and alkali zones. They reported that with Bacillus coli 
fermentative activity was checked in 1 per cent dextrose bouillon 
at P H 5.0, but that growth in plain bouillon was checked at 
P H 5.7. They noted evidence of inhibition which obscured their 
results, but they did not study the inhibitory factors; however, 
it was found that the period of lag was more pronounced in 
alkaline than in acid media. 

Recent contributions from Besson, Ranque, and Senez ('19), 
while they do not involve hydrogen ion concentration, advance 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 255 



some new ideas on sugar relations and fermentation. They 
worked with Bacillus coli in bouillon containing varying amounts 
of dextrose. With less than .4 per cent dextrose the sugar 
was all removed in 24 hours and the cultures were viable after 
10 days, while with .4 per cent or over the cultures were sterile 
in 6 days. They reported that fermentation with gas com- 
menced at the time multiplication of the organisms ceased, that 
acid production started at the same time, and that more than 
one-half the total acid is produced in 1 hour. 

From the literature reviewed it would appear that a corre- 
lation of growth curves and P H curves, with frequent observa- 
tions during growth, rather than a study of final hydrogen ion 
concentration, would add to our knowledge of metabolic changes 
in hydrogen ion concentration and of inhibition during growth. 

Technique 

The experimental work was planned on the basis of a corre- 
lation of the growth of the bacteria with the changes in the 
hydrogen ion concentration of the media produced during growth. 
The technique was uniform throughout to make all results com- 
parable. Cultures were grown in Florence flasks of 500, 1000, 
and 2000 cc. capacity, filled to one-half their capacities for the 
initial volume of media to insure a uniform and maximum sur- 
face. The basic bouillon for all the cultures, designated plain 
bouillon through the text, consisted of 2.5 per cent bacto-beef 
and 1 per cent bacto-peptone made up with distilled water 
according to the Digestive Ferments Company circular of De- 
cember, 1916. This plain bouillon forms the basis for the 
different dextrose media, with a few exceptions which are noted 

in the data. 

A culture of Bacillus coli, culture FG, kindly furnished from 
the Dairy Division, United States Department of Agriculture, 
was used throughout the experimental work with one excep- 
tion, in which Bacillus aerogenes, culture VE from the same 
laboratory, was substituted. 

To avoid the lag phase, transfers from stock agar were grown 
through two successive cultures of plain bouillon, and the inocu- 
lation was made from the second culture between 6 and 10 hours, 
during its period of logarithmic increase. A uniform tempera- 



25G 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



GROWTH AND 

PLAIN 



TABLE I 

HYDROGEN ION CONCENTRATION OF BACILLUS COLI 
AND 1 PER CENT DEXTROSE BOUILLON AT 30° C. 



IN 





1 




2 
Plain bouillon and 




Flam bouillon 


1% dextrose 


1 


Hours 






/ V* 






Bacteria per cc. 


Ph 


Bacteria per cc. 


55,000 


Pei 





- 

54,000 


7.1 


7.1 


12 


175,000,000 


6.8 


268,000,000 


5.3 


20 






281,000,000 


5.1 


24 


320,000,000 


7.2 


214,000,000 


4.9 


36 


538,000,000 


7.5 


220,000,000 


4.8 


48 


609,000,000 


7.6 


189,000,000 


4.8 


72 


450,000,000 


7.7 


119,000,000 


4.8 


96 


459,000,000 


7.8 


14,500,000 


4.9 


120 


293,000,000 


7.9 


11,100 


4.9 


144 


250,000,000 


8.1 





4.9 


168 


151,000,000 


8.1 






192 


156,000,000 


8.3 






234 


125,000,000 


8.2 






276 


115,000,000 


8.3 






348 


89,000,000 


8.3 






492 


69,000,000 


8.3 






612 


71,000,000 


8.3 






1284 


53,000,000 


8.5 






1800 


7,500,000 


8.7 







C. was maintained for all cultures throughout the 



work 



The chan 



in 



each culture were 



th and hyd 



concentration in 



ed by removing, under aseptic con- 



ditions, a 3-cc. sample at 12- or 24-hour intervals, after the flasks 
had been rotated briskly 30 times to mix the contents thor- 



oughly. One 



diluted and plated in triplicate in agar 



iposed of plain bouillon 



1 per cent dextrose and 



1.5 per cent granular agar had been added. The hydrogen 
ion concentration was determined from the remaining 2 cc. 
according to the colorimetric method of Clark and Lubs f 17). 



using the micro-colorimeter described by Dug 



('19). The 



hydrogen ion concentration is expressed in P H , or the 



r alues of Sorensen now in general biolo 
ounted after an incubation of 72 hours 



Plates were 



C 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 257 



Bacteria per cc. 
(Logs.) 



10.0 



P 



K 





Hours 







48 



96 



144 



192 



240 



Fig. 1. Growth and hydrogen ion concentration of Bacillus colt at 30° C. 

plain bouillon. 



- 1 per cent dextrose bouillon. 



[Vol. 7 



258 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Experimental Data 

As a starting point for the experimental work and as a basis 
for comparison of inhibitory action, one culture in plain bouillon 
and one culture in this bouillon with 1 per cent dextrose added 
were inoculated with equal numbers of Bacillus coli from the 
same culture. The resulting growth (expressed in numbers of 
bacteria per cc), and the hydrogen ion concentration of the 
media (expressed in P H ) are recorded in table I. The compari- 
son is more strikingly shown in fig. 1, in which the growth curves 
are plotted from the logarithms of numbers of bacteria per cc. 
as given in table i. A comparison of the hydrogen ion curves 
shows a rapid production of acid from dextrose, attaining P H 4.8 
in 36 hours, but a slower production of alkali in the plain bouillon 
with the exception of the short acid break at the beginning 
of the curve. Growth in the dextrose bouillon is more rapid in 
12 hours than in the plain bouillon but the maximum is reached 

in 20 hours, 281,000,000 bacteria per cc. when the P H is 5.1, 
and the decline is then very abrupt, terminating in sterility of 
the culture in 144 hours. In the plain bouillon, the maximum 
is reached in 48 hours, 009,000,000 bacteria per cc, with a P H 
of 7.6. However, after 75 days, although a P H of 8.7 is attained, 
there are still 7,500,000 viable bacteria per cc. in the culture. 
Apparently, then, the more intense inhibition is found in the 
dextrose rather than in the plain bouillon. 

If a bacterial "autotoxin," or any inhibitory action such 
as Chesney found with pneumococcus in plain broth, is produced 
by Bacillus coli, it would seem, from the results given in table i, 
to be associated with the dextrose bouillon and not with the 
plain bouillon. A series of cultures in a 1 per cent dextrose 
medium were observed for the purpose of determining any 
variation in inhibitory action during growth and death. The 
results are given in table n and illustrated in fig. 2. Flask 1, 
the parent culture, contained the same 1 per cent dextrose 
bouillon as Culture 2 of table l. Subcultures of 200 cc. each 
were removed from the parent cultures at the times indicated, 
commencing before the point of maximum growth was reached 
and covering a range well into the period of rapid death. The 
reaction of the subcultures was readjusted to approximately 
neutral with sterile N/1 NaOH to eliminate the acidity factor, 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 259 



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[Vol. 7 



260 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Bacteria per cc 

(Logs.) 
10.0 



PH 



9.0 



8.0 



7.0 



6.0 



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192 



240 



Fig. 2. Growth and hydrogen ion concentration of Bacillus coli at 30° C, 1 per 
cent dextrose bouillon, subcultured at intervals. 

parent culture. 

Subculture 1 . 

Subculture 2. 



• Subculture 3. 

Subculture 4. 

• • • Subculture 5. 



1 920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 20 1 



and the subcultures were then observed as new cultures. The 
points of maximum growth for the different subcultures in order 
are 557,000,000, 428,000,000, 342,000,0C0, 264,000,000, and 
274,000,000 bacteria per cc, so that the subcultures fall in a 
regular series of decreasing maximum growths, with the excep- 
tion of Culture 4 whose maximum might have occurred unob- 
served between 84 and 96 hours. There is nothing distinctive 
in the changes in hydrogen ion concentration, each subculture 
producing acid to P H 4.9 in 24 to 36 hours. It would seem, 
then, from this series of subcultures than some factor besides 
hydrogen ion concentration caused an inhibition of the growth, 
increasing with the age of the culture. 

To determine the influence of the exhaustion of the medium 
as a factor in the inhibitory action shown in fig. 2, another simi- 
lar series was observed. Four subcultures of 200 cc. each were 
removed from a parent culture at 96 hours and treated in differ- 
ent ways. All were readjusted to approximately neutral with 
N/1 NaOH. In addition 50 cc. of plain bouillon condensed 5 
times was added to No. 2 (200 cc), making a total volume of 
250 cc. No. 3 received 50 cc. of the condensed bouillon and 
1 per cent dextrose. No. 4 received the same nutrients as No. 3 
and was then sterilized for 15 minutes at 120° C. to kill any 
"autotoxin" or inhibitory enzymatic substance, and reinocu- 
lated as a new culture. The results are contained in table in 
and fig. 3. The changes in hydrogen ion concentration are 
very uniform, falling on almost the same line, an increase until 
P H 4.9 is reached in 48 hours. The exhaustion of the medium 
is shown, however, by the increased growth both in the culture 
with the bouillon replenished and in the replenished bouillon with 
dextrose added. The addition of dextrose shows almost no 
advantage over the addition of concentrated bouillon alone, 
so that dextrose is not considered an important factor at this 
time. Acid production from P H 7.3 to 4.9 in Subcultures 1 and 2 
where the dextrose was not replenished shows that all the dex- 
trose had not been fermented in the parent culture at 96 hours. 
The maximum in Subculture 3 (table in) of 545,000,000 bacteria 
per cc. compares very favorably with 5.37,000,000 in Subcul- 
ture 1 of table ii, so that it would appear that the exhaustion 
of the nutrients contained in the plain bouillon was a very im- 



262 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 






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1 920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 263 



Bacteria per cc 
(Logs.) 



10.0 



P 



K 








Hours o 



48 



96 



144 



192 



240 



Fig. 3. Growth and hydrogen ion concentration of Bacillus coli at 30° C, 1 per 
cent dextrose bouillon, subcultures with added nutrients. 

parent culture. 



• * 



Subculture 1. 
Subculture 2. 
Subculture 3. 
Subculture 4. 



[Vol. 7 



264 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



portant factor in causing the increasing inhibitory action up 
to 96 hours. Subculture 4, with the same added nutrients as 
Subculture 3 but sterilized and reinoculated, did not attain the 
growth of Subculture 3, probably because of the small inocula- 
tion. However, the fact that the sterilized subculture did not 
surpass the unsterilized would indicate that in the parent cul- 
ture or in the other subcultures the inhibition was not due to 

■ 

a substance which could be killed by sterilizing. 

Some investigators have reported that the inhibitory action 
disappeared on standing and that a good growth was attained 
upon reinoculation, although the acidity was unaltered. To 
check this with Bacillus coli, 3 to 5 days after the cultures re- 
ported in table in became sterile, Subcultures 1, 3, and 4 were 
mixed together and divided into three equal 200-cc. portions, 
designated Cultures A, B, and C. Culture A was unchanged; 



Culture B 



C: and Culture C 



was readjusted to P H 7.3. All were inoculated from the same 
culture of Bacillus coli with approximately 275,000 bacteria 
per cc. The growth and hydrogen ion concentration changes 
are recorded in table iv and fig. 4. Where unaltered, the hy- 
drogen ion concentration in Cultures A and B is P H 5.1 at inocu- 
lation, progressing to P H 4.9 in a short time. Death of the 
bacteria occurs shortly, with very little difference between the 
sterilized and unsterilized cultures. In Culture C, unsterilized 
but with acidity corrected to P H 7.3, growth and formation 
of acid occur similar to that in a new culture. Normal growth 
when the acidity was adjusted to neutral and no growth when it 
was not, both in the sterilized and unsterilized cultures, would indi- 
cate that no thermolabile substance which disappears on stand- 
ing was present and that the hydrogen ion concentration of 

the medium was the important inhibitory factor. 

The combined results expressed in the four tables might be 
summarized as follows: Inhibition to the point of death occurred 
only in dextrose bouillon in conjunction with acid formation, 



and not in plain bouillon 



A slight 



found in dextrose bouillo 



the age of the culture up to 96 hours, which was not attributable 
to acid but which probably was due to a diminution of the 
nutrients in the medium. No indication was found of an in- 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABCL1C PRODUCTS 265 



TABLE IV 

I 

GROWTH AND HYDROGEN ION CONCENTRATION OF BACILLUS COL1 AT 

30° C. f DEXTROSE BOUILLON, RE1NOCULATED 



Culture 


A 


B 


C 


Treatment 


None 


Sterilized 


Acidity adjusted 


Hours 

> 


Bacteria 
per cc. 


Ph 


Bacteria 
per cc. 


Ph 


Bacteria 
per cc. 


Ph 



12 

24 

36 

48 

60 

84 

108 
132 

156 

180 

204 

228 
252 
300 


275,000 

11,400 

7,200 

3,600 

530 

30 




5.1 
5.0 
5.0 

4.9 
4.9 
4.9 
4.9 


286,000 

15,200 

3,800 

650 

75 

2 



• 


5.1 
4.9 
4.9 
4.9 
4.9 
4.9 
4.9 


261,000 
280,000,000 
320,000,000 
340,000,000 

264,000,000 

104,000,000 

89,000,000 

57,000,000 

9,500,000 

660,000 

36,600 

5,400 

280 




7.3 
5.5 
5.5 
5.3 

5.1 
5.1 
5.1 
5.1 
5.1 
5.1 
5.1 
5.1 
5.1 
5.1 



hibitory substance which was destroyed by sterilization or 
inactivated on standing. The evidence of these results is against 



an "autotoxin" theory and points toward the hydrogen ion 
concentration as the predominating inhibitory factor in the 

experiments cited. 

The balance of the experimental work concerns the relation 
of hydrogen ion concentration to inhibition. To counteract 
the influence of acid and alkali produced during growth, and 
thus to study their action by comparison, two cultures were 
observed in which the acid or alkali formed was neutralized at 
frequent intervals. The media used was the same as that 
reported in table l, one culture of plain bouillon and the other 
of 1 per cent dextrose bouillon, 500 cc. each in 1000-cc. flasks. 
The acid produced in the dextrose culture was neutralized at 
12-hour intervals by the addition of N/1 NaOH and the hy- 



266 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Bacteria per cc. 
(Logs. ) 

10.0 




?H 



4.0 



5.0 



6.0 



7.0 



8.0 




r 



9.0 



. 



. 



Hours 



o 



48 



96 



144 



192 



240 



Fig. 4. Growth and hydrogen ion concentration of Bacillus colt at 30° C, dex- 



trose bouillon, reinoculated. 



Culture A, untreated. 
Culture B, sterilized. 
Culture C, acidity adjusted. 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 267 



TABLE V 

GROWTH AND HYDROGEN ION CONCENTRATION OP BACILLUS COLI AT 

30° C, PLAIN AND 1 PER CENT DEXTROSE BOUILLON, 

NEUTRALIZED AT INTERVALS 



1 

Plain bouillon 



Hours 




12 
14 

24 
26 
36 
38 
48 
50 
60 
62 
72 
74 
96 



98 

120 

122 

144 

146 

168 
192 

234 

236 

276 

278 

348 

350 

492 



Bacteria 
per cc. 



57,000 
184,000,000 



391,000,000 



552,000,000 



660,000,000 



631,000,000 



684,000,000 



696,000,000 



680,000,000 



570,000,000 
693,000,000 
437,000,000 



450,000,000 



327,000,000 



256,000,000 



Cc. of 

N/5 HC1 
added 



3 



5 



5 



3 



3 



3 



3 



3 



3 



4 



3 



Ph 



7.1 
7.0 



7.3 
6.9 
7.4 
7.0 
7.3 
6.7 
7.1 
6.7 
7.1 
6.7 
7.2 



6.8 
7.3 
6.7 
7.3 
6.9 
7.2 
7.1 
7.3 
6.9 
7.1 
6.8 
7.1 
6.5 
7.1 



2 

1% Dextrose bouillon 



Bacteris 
per cc. 



i 






54,030 
259,000,000 



347,000,000 



507,000,000 



612,000,000 



692,000,000 



728,000,000 



532,000,000 



455,000,000 



487,000,000 
618,000,000 
885,000,000 



794,000,000 



608,000,000 



441,000,000 



Cc. of 

N/1 NaOH 

and N/5 HC1 

added 



N/1 NaOH 



4.0 



6.3 



6.5 



7.0 



7.0 



7.0 



N/5 HC1 
3.0 



3.0 



16.0 



15.0 



20.0 



25.0 



Ph 



7.1 
5.3 
6.5 
5.1 
7.1 
5.1 
6.9 
5.1 
7.0 
5.3 
6.7 
5.7 
7.3 
7.5 



7.3 
7.3 
7.3 
7.5 
6.9 
7.5 
7.3 
7.5 
7.0 
7.6 
7.1 
7.7 
6.7 
8.1 



drogen ion concentration was determined before and after each 
addition. The plain bouillon was treated similarly, correcting 



the alkali with N/5 HC1. The 



the 



hydrogen ion concentration and the cc. of acid or alkali added 



[Vol. 7 



2G8 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Bacteria per cc 
(Logs. ) 



10.0 



p 



H 




4.0 



5.0 



6.0 



7.0 



8.0 



9.0 




Hour 8 



O 



48 



96 



144 



192 



240 



Kig. 5. Growth and hydrogen ion concentration of Bacillus coli at 30° C, plain 
and 1 per cent dextrose bouillon, neutralized at intervals. 



plain bouillon. 

1 per cent dextrose bouillon. 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 269 



are given in table v. A more striking representation of the 
changes in the concentration of the hydrogen ions is shown 
in the curves in fig. 5. Growth in both cultures is practically 
parallel, with the dextrose culture reaching the highest point — 
885,000,000 per cc. at 234 hours. From a comparison of these 
growth curves with those of fig. 1, it is quite evident that neu- 
tralizing the acid or alkali prolongs the growth at a higher level. 
The P H curve for the dextrose culture shows an abundant 
production of acid, going as high as P H 5.1 several times. Be- 
tween 72 and 96 hours, however, the formation changed to alkali, 
and N/5 HC1 was added to neutralize. Table v shows that 
37.8 cc. of N/1 NaOH were required to neutralize the acid 
from 1 per cent dextrose and that in the same time, 96 hours, 
19 cc. of N/5 HC1 were used in neutralizing the alkali in the 
plain bouillon, giving a ratio of 189 to 19, or approximately 

10 to 1. Theoretically, then, one-tenth of the dextrose, or .1 
per cent dextrose, would furnish just enough acid in 96 hours to 

neutralize the alkali formed in plain bouillon, and would hold 
at neutral the hydrogen ion concentration of a growing culture 
which was fermenting dextrose, if the dextrose were added in 
small amounts at frequent intervals. 

On this basis a culture was started in plain bouillon. The 
amounts of dextrose added, the growth, and the P H values are 
given in table vi and illustrated in fig. 6. By 72 hours the 
hydrogen ion concentration had demonstrated that the the- 
oretical amount, .025 per cent of dextrose every 24 hours, did 
not furnish sufficient acid to neutralize the alkali, so the amount 
of dextrose was increased and the intervals between additions 
shortened to meet the needs of the culture. The reaction, with 
each addition of sugar, depends on the arid fermentation of 
the sugar and a subsequent alkali formation, as illustrated by 
the P H curves between 48 and 72 hours and between 96 and 108 
hours. This alkali formation was reversed by the addition of 
more sugar at the proper time. Although the theoretical cal- 
culation was upset by the increased growth, the P H curve dem- 
onstrates that it was possible to hold the hydrogen ion con- 
centration within a very narrow zone around the neutral point. 
The growth was very rapid, reaching 1,550,000,000 bacteria per 
cc. at 48 hours, or 2V£ times as many bacteria as Culture 2, 



270 



[Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



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1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 271 



Bacteria per ce. 

(Logs.) 

10.0 



9.0 



6.0 



7.0 



6.0 



5.0 




P 



H 



4.0 



5.0 



6.0 



7.0 



8.0 



9.0 




Hours 







48 



96 



144 



192 



240 



Fig. 6. Growth and hydrogen ion concentration of Banllus coli at 30° C, dex- 
trose added at intervals. 



272 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



table v, produced in the same period of growth. Probably 



this increased growth explains the more rapid utilization of 

the dextrose than was calculated. The maximum growth was 
attained at 168 hours — 3,750,000,000 bacteria per cc. Thus 
this culture showed the least inhibition of any of the experi- 
mental cultures and serves as a standard for comparison with 
the others. 

To study more in detail the effect of small amounts of acid, 
a series of cultures was observed in which the only individual 
variation was in the initial amount of dextrose. To 250 cc. 
of plain bouillon in each of five 500-cc. flasks were added re- 
spectively .05, .1, .15, .2, and .3 per cent of dextrose, and all 
were inoculated from the same culture tube of Bacillus coli. 
The growth and changes in P H are presented in table vn. The 
cultures are numbered, as indicated in the table, from 1 to 5 
in order of increasing amounts of dextrose. Cultures 2, 4, and 
5 are plotted in fig. 7 as representative of the series. The P H 
curves show that acid was produced in each culture and that 
the amount of acid formed corresponded to the amount of dex- 
trose provided. The cultures formed a regular series of increas- 
ing acidities. Following the acid production there was a rever- 
sion of the reaction toward alkalinity which was quite rapid 
in the first four cultures but slower in Culture 5, where a P 
of 5.1 was maintained from 24 to 96 hours. Comparing the 
growth curves of the five cultures during the period from inocu- 
lation to 48 hours, it is seen that Culture 2 makes the best growth 
and that Cultures 3, 4, and 5 follow in order. It would appear, 
then, that .1 per cent of dextrose or less is stimulative in effect 
and that there is no acid injury from a short exposure to P, t 5.9 
(Culture 2). There is, however, some acid inhibition from a 
P H of 5.5 (Cultures 3 and 4) and quite a marked inhibition 
sufficient to cause some decrease in numbers — from 3 days' 
exposure to P H 5.1 (Culture 5). In each case the growth curve 
ascended as the P H curve descended toward the alkaline side. 
The maximum growth was approximately the same for all the 
cultures— 1,400,000,000 to 1,800,000,000 bacteria per cc— and 
was reached when the hydrogen ion concentration fell in a zone 
between P H 7.0 and 7.6. At the point of maximum growth 
the hydrogen ion concentrations for the cultures in order were 



H 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 273 




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(Vol. 7 



274 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Bacteria per cc. 

(Logs.) 

10.0 



>H 



9.0 



8.0 



7.0 



6.0 



5.0 




4.0 



5.0 



6.0 



T.O 



e.o 



9.0 




Hours 







48 



96 



144 



192 



240 



Fig. 7. Growth and hydrogen ion concentration of Bacillus coli at 30° C, varia- 



tion in initial dextrose. 



.1 per cent dextrose. 
.2 per cent dextrose. 
.3 per cent dextrose. 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 275 



P H 7.3, 7.1, 7.6, 7.5, and 7.4. The relation between growth 
curves and P H curves, fig. 7, would indicate that Bacillus coli 
is more sensitive to alkali than to acid and that amounts of 
alkali or acid considerably less than the fatal dose become promi- 
nent factors in inhibiting growth. 

Supplementing the preceding table, table vni gives the results 
of growth of Bacillus coli in 1 per cent, 2.5 per cent, and 5 per 
cent dextrose media. A synthetic bouillon was used for these 
cultures consisting of .5 per cent asparagin, .5 per cent K 2 HP0 4 , 
and the dextrose as indicated. The growth and hydrogen ion 
concentration curves are plotted in fig. 8 on the same basis as 
the curves in all the other figures. As might be expected, the 
action in general corresponded to that of Culture 2, fig. 1, which 
was grown in 1 per cent dextrose. Both the growth and P„ 
curves showed a small lag at the beginning in 2.5 per cent dex- 
trose and a greater one with some decrease in growth in 5 per 
cent dextrose. Following the initial lag, the cultures produced 
the usual growth, acid fermentation, and death. A slightly 
greater acid production occurred in the 5 per cent dextrose, 
for the hydrogen ion concentration went to P H 4.7. The data of 
tables vn and vm show that in cultures of Bacillus coli sufficient 
acid to kill the organisms was formed from 1 per cent or more 
of dextrose, while .15 to .3 per cent supplied only enough acid 
to inhibit the growth, and .1 per cent exerted a stimulative 
action. Thus the amount of dextrose present seems to regu- 
late the reaction, which is a strong factor in growth and inhi- 
bition. 

In connection with the reversion of reaction, the growth 
and inhibition of Bacillus aerogenes are of interest. One culture 
of plain bouillon and one culture of plain bouillon plus 1 per 
cent dextrose were inoculated with Bacillus aerogenes; the growth 
and hydrogen ion concentration changes are recorded in table ix 
and fig. 9. Both the growth and hydrogen ion concentration 
were very similar to those of Bacillus coli from the time of inocu- 
lation up to 96 hours. As fig. 9 illustrates, at 96 hours the abrupt 
descent of the growth curve was checked at 7,600,000 bacteria 
per cc. The slight drop to 5,000,000 bacteria per cc. in the 
next 48 hours was followed by a second rise which culminated in 
a maximum of 1,017,000,000 bacteria per cc. at 696 hours. 



276 



(Vol. 7 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



TABLE VIII 

GROWTH AND HYDROGEN ION CONCENTRATION OP BACILLUS COL1 AT 

30° C, VARIATION IN INITIAL DEXTROSE IN 

ASPARAGIN BOUILLON 



Cultures 


1 


2 




3 


Dextrose 


10' 


r 

) 


2.5% 

1 


) 


5.0% 


Hours 


Bacteria 


Ph 


Bacteria 


Ph 


Bacteria 


Ph 




per cc. 




per cc. 




per cc. 







45,000 


6.6 


63,000 


6.4 


46,000 


6.4 


12 


5,450,000 


6.6 


100,000 


6.4 


60,000 


6.3 


16 


46,500,000 


6.5 


80,000 


6.3 


60,000 


6.2 


20 


326,000,000 


5.7 


80,000 


5.8 


60,000 


5.9 


24 


350,000,000 


5.5 


145,000 


5.7 


25,200 


5.9 


36 


464,000,000 


5.4 


17,000,000 


5.8 


8,400 


5.9 


48 


416,000,000 


4.9 


345,000,000 


5.6 


57,600 


5.8 


60 


280,000,000 


4.9 


384,000,000 


4.9 


1,700,000 


5.7 


72 


178,000,000 


5.3 


323,000,000 


4.9 


40,000,000 


5.7 


84 


108,000,000 


5.3 


348,000,000 


4.9 


170,000,000 


5.1 


96 


51,000,000 


5.3 


320,000,000 


4.9 


270,000,000 


4.9 


108 


28,000,000 


5.3 


180,000,000 


4.9 


260,000,000 


4.9 


120 


1,800,000 


5.3 


187,000,000 


4.9 


380,000,000 


4.7 


132 


2,800,000 


5.3 


120,000,000 


4.9 


224,000,000 


4.7 


144 


200,000 




31,000,000 




142,000,000 


4.7 


156 


18,400 


5.3 


24,000,000 


4.9 


161,000,000 


4.9 


168 


1,000 


5.3 


6,000,000 


5.2 


158,000,000 


5.1 


180 


346 


5.3 


2,000,000 


5.2 


132,000,000 


5.1 


192 


144 


5.3 


360,000 


5.0 


102,000,000 


4.9 


204 


93 




62,000 




104,000,000 




216 






391 


5.2 


54,000,000 


5.3 


240 










21,000,000 


5.3 


264 










9,900,000 


5.0 


300 










900 


5.0 



Some points of interest in the P H curves are that the high point, 
P H 4.7, at 120 hours, did not occur during the period of greatest 
decrease in growth, that the P H held at 4.9 for a considerable 
period after the second increase in growth began, and that 
the hydrogen ion concentration at the time when the maximum 
growth was reached was P H 7.1. In plain bouillon there was 
no essential difference in growth or changes in hydrogen ion 



12901 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 277 



Bacteria per cc 

(Logs. ) 

9.0 



P 



H 



8.0 



7.0 



6.0 



5.0 



4.0 



4.0 



5.0 



6.0 



7.0 



8.0 



9.0 



Hours 







48 



96 



144 



192 





840 



Fig. 8. Growth and hydrogen ion concentration of Bacillus coli at 30° C, varia- 
tion in initial dextrose in asparagin bouillon. 



1.0 per cent dextrose. 
2.5 per cent dextrose. 
5.0 per cent dextrose. 



|Vol. 7 



278 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



TABLE IX 

GROWTH AND HYDROGEN ION CONCENTRATION OF BACILLUS AERO 
GENES AT 30° O., PLAIN AND 1 PER CENT DEXTROSE BOUILLON 



Hours 




12 

24 

48 

72 

96 

120 

144 

168 

19 

216 

264 

312 

360 

408 

456 

504 

552 

624 

696 

864 

1032 

1200 



1 
Plain bouillon 



Bacteria per cc. 



98,000 
171,000,000 
236,000,000 
306,000,000 
455,000,000 
307,000,000 
169,000,000 



162,000,000 
121,000,000 
108,000,000 



37,000,000 
44,000,000 



36,000,000 

20,000,000 

22,000,000 

8,640,000 

2,520,000 

7,830,000 



P.. 



0.9 
6.9 
7.0 

7.4 
7.8 
8.1 
8.3 
8.3 
8.3 
8.3 
8.3 



8.3 
8.5 



8.6 
8.7 
8.7 
8.7 
8.7 
8.7 



2 

Plain bouillon and 

1 % dextrose 



Bacteria per cc. 



P.. 



94,000 

272,000,000 

300,000,000 

139,000,000 

27,000,000 

7,600,000 

6,300,000 

5,000,000 

10,000,000 

23,000,000 

38,000,000 

125,000,000 

211,000,000 

539,000,000 

815,000,000 

836,000,000 

761,000,000 

775,000,000 

927,000,000 

1,017,000,000 

690,000,000 

643,000,000 

485,000,000 



6.9 

5 . 5 

4.9 

4.8 

4.8 

4.8 

4.7 

4.9 

4.9 
4.9 

4.9 
4.9 
4.9 
5.8 
6.6 
6.9 
6.9 
6.9 
6.9 
7.1 
7.3 
7.0 

7.6 






concentration betw 



Bacillus coli (table I, Culture 




and 



Bacillus 



(table 



Culture 1). There 



growth of Bacillus coli in .3 per cent dextrose and B 



dextrose, the difference betw 



ganisms 



being 



of Ba 



acid 



While the experimental data reported above have emphasized 
e imoortance of the H and OH ions as factors in inhibition, 



these ions do not represent the 



products of metabolism 



hich might be considered as inhibitory to growth. Ayers and 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 279 



Bacteria per* cc. 

10.0 



p 



H 



9.0 



8.0 



7,0 



6.0 



5.0 




4.0 



5.0 



6.0 



7.0 



8.0 



9.0 




Hours 







192 



384 



576 



768 



960 



Fig. 9. Growth and hydrogen ion concentration of Bacillus aerogenes at 30° C. 

plain bouillon. 



1 per cent dextrose bouillon. 



[Vol. 7 



280 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Rupp have made quantitative determinations of formic, acetic, 
lactic, and succinic acids from Bacillus coli in a dextrose bouillon, 
and Wyeth, and Cohen and Clark have shown that the critical 
hydrogen ion concentration varies with the different acids, 
hydrochloric, acetic, and lactic, indicating that the anions of 
the acids or perhaps the undissociated molecules, as Winslow 
and Lochridge suggested, are also concerned in inhibition. Most 
of the work on the inhibitory effect of different acids has been 
based on inoculation of media of different P H values obtained 
by using different acids, and the inhibition has been determined 
according to the presence or absence of growth after a certain 
interval. Such a method does not take into consideration 
milder phases of inhibition which are not severe enough to cause 
the death of the organisms. To illustrate this phase and to 
indicate some of the relations of the hydrogen ion factor to 
the other factors, the results of an experiment are presented in 
table x and fig. 10. Culture 1 was grown in 1 per cent dextrose 
bouillon, and Culture 2 in plain bouillon to which sterile N/5 
HC1 was added, as indicated in table x, in an attempt to simu- 
late in plain bouillon the P H curve of a culture fermenting dex- 
trose bouillon, such as Culture 1. As seen from fig. 10, the 
culture produced alkali continually so that it was only possible 
by frequent additions of acid to hold the P H in a zone around 
P H 4.8, the greatest hydrogen ion concentration which Culture 1 
attained. The growth curve, fig. 10, shows marked acid inhibi- 
tion with almost no further increase in growth after the first 
addition of acid at 14 hours. There is practically no difference 



the growth curves of 



but 



from that point they separate widely, for death occurs shortly 
in the dextrose media and growth in Culture 2 does not go below 
26,000,000 bacteria per cc. Thus a hydrogen ion concentra- 
tion of P H 4.8-4.9 when produced by the acid fermentation of 
dextrose was fatal, while that of P H 4.7-5.1 from HC1 was 
only strongly inhibitory, indicating that the other metabolic 
products of dextrose fermentation, such as acetate or lactate 
ions, evidently enter as factors in causing the death of the culture. 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 281 



TABLE X 

GROWTH AND HYDROGEN ION CONCENTRATION OF BACILLUS COLI AT 

30° C, 1 PER CENT DEXTROSE BOUILLON AND PLAIN 

BOUILLON +HC1 AT INTERVALS 



Hours 





12 

14 

20 
24 

26 

36 

38 

48 

50 

60 

72 

74 

96 

120 

122 

144 

146 

168 

192 

194 

234 

276 
278 
348 
350 
492 
612 



1 

1% dextrose bouillon 



Bacteria 
per cc. 



55,000 
268,000,000 



281,000,000 
214,000,000 



220,000,000 



189,000,000 



119,000,000 



14,500,000 
11,100 







Ph 



7.1 
5.3 



5.1 

4.9 



4.8 



4.8 



4.8 



4.9 
4.9 



4.9 



2 
Plain bouillon + HC1 



Bacteria 
per cc. 



57,000 
179,000,000 



178,000,000 



197,000,000 



175,000,000 



116,000,000 



77,000,000 
96,000,000 



55,000,000 



26,000,000 
60,000,000 



74,000,000 
144,000,000 



148,000,000 



95,000,000 
73,000,000 



Cc. of 

N/5 HC1 
added 



15 



4 



5 



5 



3 



3 



2 



2 



4 



3 



P 



ii 



7.1 

6.7 
5.2 



5.3 
5.0 
5.2 
4.9 
5.1 
4.7 
4.9 
5.0 
4.7 
4.9 
5.3 
4.9 
5.1 
4.9 
4.9 
5.1 
4.9 
4.9 

5.3 
4.9 
5.1 
4.8 
4.9 
5.1 



Discussion 

For a discussion of the combined results embodied in the 
experimental data presented, the inhibitory products of meta- 
bolism of Bacillus coli are divided under four topics: "auto- 
toxins"; H ions; OH ions; and products of dextrose fermenta- 
tion other than those directly related to changes in H ion con- 
centration. 



■Vol. 7 



282 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Bacteria bar e«. 

(Logs. ) 

10.0 



P 



H 




4.0 



5.0 



6.0 



T.O 



8.0 



9.0 




Hours 







46 



96 



144 



192 



240 



Fig. 10. Growth and hydrogen ion concentration of Bacillus coli at 30° C. 

plain bouillon +HC1 at intervals. 

1 per cent dextrose bouillon. 



1920] 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 283 



It appears improbable that Bacillus coli, grown under the 
conditions of the experiments reported, produces any ' 'auto- 
toxin" or special inhibitory substance such as Eijkmann and 
others claimed. The results given in tables n and in indicate 
that the slight inhibition increasing with the age of the culture 
is probably associated with a diminution in nutrients. No 
thermolabile product could be detected wiich would inhibit 
growth if the hydrogen ion concentration were corrected. It is 
difficult to reconcile the production of an enzymatic inhibiting 
substance with such growth as appears in figs. 1 and 6, especially 
as death did not occur in these cultures. In plain bouillon, 
fig. 1, the culture was viable after 75 days; and in dextrose 
bouillon, fig. 6, with the hydrogen ion factor controlled, growth 
attained 3,750,000,000 bacteria per cc. in 7 days, and there 
were still present over 2,000,000,000 bacteria per cc. after 840 
hours. In addition there was no indication in the cultures in 
which death occurred that death could be attributed to an 
"autotoxin." 

There is a direct relation between hydrogen ion concentration 
and inhibition. If the acid is formed from the fermentation of 
dextrose, with Bacillus coli, fig. 7 and table vn, there is no indi- 
cation of acid inhibition at P H 5.9 if maintained for only a short 
time. Some inhibition is apparent at P H 5.5 which increases 
with the time that the culture is exposed to this P H . There 
is a marked inhibition from an exposure of 72 hours to P H 5.1, 



fig. 7, but it is insufficient to cause death. Fig. 4, however, 



shows that a prolonged hydrogen ion concentration of P H 5.1 
is lethal, and in every case throughout the experimental work 
a P H of 4.9, when produced by acid fermentation of dextrose, 
proves fatal, figs. 1, 2, 3, 4, and 8. 

To illustrate the relationship between hydrogen ion concen- 
tration and growth, four curves from figs. 1, 6, and 7 are as- 
sembled in fig. 11. The highest growth curve, No. 2, is attained 
in the culture in which the P H remains practically neutral; the 
P H of 5.1, No. 3, produces an intermediate growth; and the slight 
difference in hydrogen ion concentration between P H 5.1, No. 3, 
and 4.9, No. 4, is fatal. 

The OH ions also prove to be inhibitory according to the 
plain bouillon growth curve in fig. 11. An alkalinity corre- 



IVol. 7 



284 



ANNALS OF THE MISSOUM BOTANICAL GARDEN 



Bacteria per 

(Logs,) 
10.0 



cc 




P 



H 



4.0 



5.0 



6.0 



7.0 



8.0 



9.0 




Hours 



48 



96 



144 



192 



240 



Fig. 11. Relation of growth to hydrogen ion concentration of Bacillus coli at 



30° C. 



plain bouillon. 



No. 1 

No. 2 - - dextrose added at intervals 

No. 3 

No. 4 



.3 per cent dextrose. 
1 per cent dextrose. 



1920 J 



CHAMBERS — BACTERIAL INHIBITION BY METABOLIC PRODUCTS 285 



sponding to P H 7.6-7.8 is comparable in toxicity with an acidity 
of P H 5.1. Bacillus coli seems more sensitive to small amounts 
of alkali than to small amounts of acid, for in the reversions 
of reaction in fig. 7, inhibition is evident shortly after crossing 
the neutral line, about P H 7.1-7.6. In a freshly inoculated 
culture without dextrose, fig. 7, inhibition is first noted about 
P H 7.5. While the hydroxyl ions appear mere toxic to Bacillus 
coli in less concentration than the hydrogen ions, they do not 
seem to be fatal in greater concentration, for death of the cul- 
ture was not observed on the alkaline side, although one culture 
containing CaC0 3 , which is not reported in the data, carried 
the P H to 9.5. 

The importance of the factors other than H or OH ions which 
may enter into the inhibition or killing of a culture of Bacillus 
coli is not overlooked, but it should not be over-emphasized. 
For example, in a 1 per cent dextrose bouillon culture, such as 
is shown in fig. 1 or 10, in addition to the H ions, the anions, 
formate, acetate, lactate, and succinate, are formed (Ayers and 
Rupp, '18), probably other anions, and also the undissociated 
acids. These add their inhibitory action to that of the H ions in 
producing death at P H 4.9, illustrated by fig. 10. The best growth 
curve, however, fig. 11, has only the hydrogen ion concentration 
controlled and in reality ferments much more dextrose than 
the 1 per cent dextrose culture of fig. 10. The former culture 
ferments a total of over 1.36 per cent and none of the products 
are removed from the culture, while the latter does not ferment 
all of the 1 per cent dextrose furnished. It seems possible that 
the metabolic products other than the H ions are not sufficiently 
inhibitory to influence greatly the growth until the hydrogen 
ion concentration approaches the acid limit, but toward the 
critical acid zone their effect becomes noticeable. 

The growth curves as a whole do not agiee exactly with the 
life phases presented by Buchanan ('18). In fact, the diversity 
of growths produced by varying the hydrogen ion concentra- 
tion, as illustrated by fig. 11, is so great that one curve can 
express the growth of Bacillus coli in bouillon only when quite 
definite limitations of conditions are imposed. In a growing 
culture of an organism like Bacillus coli which produces acid 
from dextrose and alkali in plain bouillon, growth can be con- 



[Vol. 7 



286 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



trolled to a certain extent by the hydrogen ion concentration, 
which can in turn be controlled by the amount of dextrose fur- 
nished. The initial amount of dextrose determines the amount 
of acid produced or the maximum hydrogen ion concentration 

attained. The work of Clark and Lubs, Besson, Ranque and 
Senez, and the experimental data presented here give a rather 
definite idea of the action of Bacillus coli according to the amount 
of dextrose in the medium. With .3 per cent or less of dex- 
trose, insufficient acid is produced to kill the organisms; .4 per 
cent or more is sufficient dextrose to produce acid to P n 4.9 or 
better, and the culture becomes sterile in 6 days or less. An 
amount of dextrose not accurately determined, but between 
.3 and .4 per cent, probably depending to some extent on the 
buffer in the medium, should produce just enough acid, between 
P H 5.1 and P H 4.9, depending on the time of exposure, to kill 
the culture. If insufficient acid to kill the culture is produced, 
as from .3 per cent or less of dextrose, a reversion of reaction 
takes place, which Ayers and Rupp have explained with Bacillus 
aerogenes as the formation of alkaline carbonates from the or- 
ganic acids, especially from the formic and acetic acids. There 
is a similarity in reaction and in growth curves between Bacillus 
aerogenes and Bacillus coli y the main difference appearing to 
be in the greater acid resistance of Bacillus aerogenes. Growth 
in the cultures where reversion of reaction takes place seems 
to be typical. One-tenth per cent of dextrose provides a stimu- 
lation to growth, but greater amounts produce some evidence 
of acid inhibition, followed by an increase in growth with the 
reversion of the reaction and alkaline inhibition between P H 7.0 
and 7.6. The least inhibition is found in a culture in which the 
hydrogen ion concentration is held in a narrow zone around 
the neutral point — probably P H 6.0-7.0 is the best — by adding 
small amounts of dextrose at frequent intervals. Thus, with 
Bacillus coli, hydrogen ion concentration and growth within 
limits can be manipulated by the dextrose furnished. The 
growth curves emphasize not only the value of the initial reac- 
tion and composition of the medium, but also the importance 
in physiological studies of following the changes in hydrogen 
ion concentration which the growing bacteria produce in their 
substrates. 



1920] 

CHAMBERS — BACTERIAL INHIBITION BY METAEOLIC PRODUCTS 287 



Summary 

Growth and death of Bacillus coli in the culture bouillon of 
these experiments does not follow a constant curve but is de- 
pendent on the hydrogen ion concentration of the medium. 

The hydrogen ion concentration of a growing culture of Bacillus 
coli is controlled by the composition of the medium, and par- 
ticularly by the amount of fermentable carbohydrate present. 

The maximum count, determined by the plate method, in the 
culture with the hydrogen ion concentration controlled is 3,750,- 
000,000 bacteria per cc, as contrasted with a maximum of 
281,000,000 bacteria per cc. in the 1 per cent dextrose bouillon 
with the hydrogen ion concentration uncontrolled. 

No investigation was made of the limiting influence of other 
factors, such as aeration, on the maximum number of bacteria 
per cc. in the culture where the hydrogen ion concentration 

was controlled. 
No metabolic product of the nature of a a "autotoxin" could 

be found. 

Of the products of metabolism, acid is :he most inhibitory, 
checking growth slightly at P H 5.5 and increasing in intensity 
to a lethal concentration between P H 5.1 and 4.9. 

The first inhibition on the alkaline side is noted between 
P H 7.0 and 7.6, depending on the age of the culture and other 
factors. P H 7.6 is comparable in inhibitory action to P H 5.1. 
In an asparagin-CaC0 3 bouillon, P H 9.5 is not fatal. 

The inhibitory action of the metabolic products of dextrose 
other than the hydrogen ions is only evident near the critical 
acid concentration. 



To Dr. B. M. Duggar the writer is deeply indebted. His 
aid and suggestions throughout the investigation have been 
invaluable. 



Bibliography 

Avery, O. T., and Cullen, G. E. ('19). Use of final P H in differentiation of Strep- 
tococcus hemolyticus. Jour. Exp. Med. 29: 215-233. 1919. 

p , ('19 a ). Hydrogen ion concentration of cultures of pneu- 

raococci of the different types in carbohydrate media. Ibid. 30: 359-378. 
1919. 



[Vol. 7 

288 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Ayers, S. H., Johnson, W. T., Jr., and Davis, B. J. (18). Thermal death point 
and final hydrogen ion concentration of streptococci. Jour. Infect. Dis. 23: 
290-300. 1918. 

, and Rupp, P. ('18). Simultaneous acid and alkali bacterial fermen- 



tation from dextrose and salts of organic acids. Ibid. 188-216. 1918. 
Besson, A., Ranque, A., and Senez, C. ('19). Sur la vie du colibacille en milieu 



liquide glucose. 



pt. Rend. 82: 76-78. 1919. 

, ( , 19 a ). Sur la vie des microbes dans les 



milieux liquides sucres. Ibid. 107-109. 1919. 



. — . — p 9 ^ ('19 b ). Sur la vie du colibacille en milieu 

liquide glucose. Importance des doses de glucose. Ibid. 164-166. 1919. 

Buchanan, R. E. ('18). Life phases in a bacterial culture. Jour. Infect. Dis. 23: 
109-125. 1918. 

Bunker, J. W. M. ('19). Studies of the diphtheria bacillus in culture. Jour. Bact, 
4:379-409. 1919. 

Chesney, A. M. ('16). Latent period in growth of bacteria. Jour. Exp. Med. 24- 
387-419. 1916. 

Clark, W. M. ('15). Final hydrogen ion concentration of cultures of Bacillus 
coli. Jour. Biol. Chem. 22: 87-98. 1915. 

, and Lubs, H. A. ('15). Differentiation of bacteria of the colon-aero- 
genes family by the use of indicators. Jour. Infect. Dis. 17: 160-173. 1915. 



1 f ('17). The colorimetric determination of hydrogen ion 

concentration and its application in bacteriology. Jour. Bact. 2: 1-34, 109- 
136, 191-236. 1917. 

Cohen, B., and Clark, W. M. ('19). Growth at different hydrogen ion concen- 
trations. Ibid. 4: 409-427. 1919. 

Conradi, H., and Kurpjuweit, O. ('05). Ueber spontane Wachstumshemmun^ 
der bakterien infolge Selbstvergiftung. Munch. Med. Wochenschr. 37: 1761- 
1764. 1905. 



y 1 C05 a ). Ueber die Bedeutung der bakteriellen Hem- 

mungsstoffe fur die Physiologie und Pathologie des Darmes. Ibid. 45: 2104- 
2168; 46: 2228-2232. 1905. 

Coplans, M. (W). On some vital properties of milk. Lancet 1907: 1074-1080 
1907. 

Cullen, G. E., and Chesney, A. M. ('18). Production of acid by pneumococci. 
Jour. Exp. Med. 28: 289-296. 1918. 

Duggar, B. M. ( T 19). The micro-colorimeter in the indicator method of hydrogen 
ion determination. Ann. Mo. Bot. Gard. 6: 179-181. 1919. 

Eijkmann ('04). Ueber thermolabile Stoffwechselprodukte als Ursache der natiir- 
lichen Wachstumshemmung der Mikroorganismen. Centralbl. f. Bakt 1 
Orig. 37: 436-449. 1904. 

, ('06). Ueber natiirliche Wachstumshemmung der Bakterien. Ibid. 



41 : 367-369, 471-474. 1906. 

Fred, E. B., and Loomis, N. E. ('17). Influence of hydrogen-ion concentration 
of medium on the reproduction of alfalfa bacteria. Jour. Bact. 2: 629-633 
1917. 



1 920 j 



CHAMBERS — BACTERIAL INHIBITION BY META 30LIC PRODUCTS 289 



Gillespie, L. J. ('18). Growth of potato scab organism at various hydrogen ion 
concentrations as related to comparative freedom of acid soils from potato 
scab. Phytopath. 8: 257-269. 1918. 

Itano, A. ('J 6). I. Relation of hydrogen ion concentration of media to the pro- 
teolytic activity of Bacillus subtilis. II. Proteolysis of Streptococcus ery- 
sipelatis and Streptococcus lacticus compared under different hydrogen ion 
concentrations. Mass. Agr. Exp. Sta. Bui. 167. K16. 

Kruse, W. ('10). Allgemeine mikrobiologie. pp. 156-1'H). Leipzig, 1910. 

Lord, F. T., and Nye, R. N. ('19). The relation of the Dneumococcus to hydrogen 
ion concentration, acid death-point, and dissolution of the organism. Jour. 
Exp. Med. 30: 389-399. 1919. 

Manteufel ('07). Das Problem der Entwicklungshemmung im Bakterienkulturen 
und seine Beziehungen zu den Absterbeerscheinungcn der Bakterien im Darm- 
kanal. Zeitschr. f. Hyg. 57: 337-354. 1907. 

Michaelis, L. ('14). Die Wasserstoffionenkonzentration. pp. 112-115. Berlin, 
1914. 

* 

Passini ('06). Die bakteriellen Hemmungsstoffe Conrulis und ihr Einfluss auf 



Wachstum 



Wochenschr 



1906. 



W. J. ('14). Nature of bacterial lag. Jour. Hyg. 14: 215-241. 1914. 



Rahn, Otto ('06). 



Wachst 



der Bakterien. Centralbl. f. Bakt. II. 16: 417-429 609-617. 1906. 

Roily ('06). Experimentelle Untersuchungen liber das biologische Verhalten der 
Bakterien im Dickdarm. Deutsche Med. Wochenschr. 43: 1733-1737. 1906. 

Salter, R. C. ('19). Rate of growth of Bacillus coli. Jour, Infect. Dis. 24: 260- 
284. 1919. 

Shohl, A. T., and Janney, J. H. ('17). The growth of Bacillus coli in urine at vary- 
ing hydrogen ion concentrations. Jour, of Urol. 1: 211-229. 1917. 

Winslow, C. E. A., and Lochridge, E. E. ('06). Toxic effect of certain acids upon 
typhoid and colon bacilli in relation to the degree of their dissociation. Jour. 
Infect. Dis. 3: 547-571. 1906. 

Wyeth, F. J. S. ('18). The effect of acids on the growth Df Bacillus coli. Biochem. 
Jour. 12:382-401. 1918. 

('19). The effects of acids, alkalies, and sugars on the growth and 



indole formation of Bacillus coli. Ibid. 13: 10-24. 1919. 



THE NUTRITIVE VALUE OF THE FOOD RESERVE IN 

COTYLEDONS 

B. M. DUGGAR 

Physiologist to the Missouri Botanical Garden, in Charge of Graduate Laboratory, 

Professor of Plant Physiology in the Henry Shaw School of Botany of 

Washington University 

Considerable work has been done in respect to determining 
the capacity for growth of immature and mature seed-plant 
embryos separated from the endosperm or from the cotyledons. 
Yet this work seems to have been of comparatively little sig- 
nificance in ascertaining whether, under any conditions of ger- 
mination and growth, these natural food reserves may be par- 
tially or completely substituted for in establishing the seedling 
with normal vigor in the soil or in the ust.al culture solutions. 
Reference will be made later to some of the more important 
literature bearing on the questions to be presented in this paper. 
The data here reported are, however, preliminary and intended 
primarily to give the results of some experiments (1) demon- 
strating, in those cases where the cotyledons serve as a food 
reserve, the striking importance of these seed-leaves in com- 
parison with certain organic substances as a source of food for 
the normal and vigorous establishment of the young plant under 
cultural conditions , and (2) suggesting the possibility that carbo- 
hydrate or hydrocarbon food material stored outside the em- 
bryo, as in the case of corn, may be of far less significance. 

Doubtless the assumption has been quite generally made that 
in the case of peas, beans, and other plants in which the coty- 
ledons furnish practically the entire food reserve these seed- 
leaves may constitute the chief source of organic food until the 
first green leaves are developed. It has seemed to the writer 
that interesting physiological problems might be approached 
through a critical study of the early food reserves, and pre- 
liminary tests with Canada field peas confirmed this assumption. 
Accordingly, the first series of experiments with Canada field 
peas and with field corn were made merely to determine quan- 
titatively the extent to which the excision of the cotyledons, 
or of the food supply stored outside of the embryo and the 
scutellum, influenced normal growth. 



Ann. Mo. Bot. Gard., Vol. 7, 1920 



(291) 



[Vol. 7 



292 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



In the case of the Canada field peas the seed were germinated 
on paraffined wire mesh over tap water, and growth was per- 
mitted to proceed in diffuse light until the plumules were well 
established with unfolding green leaves. Solution cultures were 
then made in the usual way as especially described in an earlier 
paper (Duggar, '19). All cultures were therefore arranged in 
duplicate in tumblers holding about 250 cc, and the seedlings 
were inserted through holes in paraffined paper covers (peas), 
or through notches in the corks (corn). Seedlings of uniform 
size were selected and all cultures were placed in the green- 
house, freely and equally exposed to sunlight. A mineral nutri- 
ent solution, designated in the paper referred to above as solu- 
tion B, was employed. It should be observed that this solution 
contains not merely all essential ions, including N0 3 , but con- 
tains these in favorable proportions and concentrations for the 
promotion of excellent growth. The date of the beginning of 
the experiment was taken as that on which the cultures were 
exposed in the greenhouse. At intervals of a day or more apart, 
the cotyledons of successive pairs of cultures were cut away 
so as to determine their influence on growth, and the time of 
excision of the last pair represented the practical exhaustion 
of these food reserves. In the case of corn the young pi ant let 
with attached scutellum was carefully dissected out from the 
endosperm, an operation which may be effected with very little 
difficulty after germination begins. In all other respects the 
corn cultures were treated in precisely the same manner as the 
peas. The total green weights of all cultures are given in table i, 
and the appearance of the peas at the end of the period of obser- 
vation, 24 days, is shown by pi. 7. 

From the results with peas it is clear that for a growth interval 
of 24 days the removal of the cotyledons after the second day 
induces a marked depression in the growth rate, and this de- 
pression is increasingly less, until, when the removal of the 
cotyledons occurs after 7 days, the amount of growth is very 
nearly the same as in the control, with cotyledons intact. Du- 
plication of this experiment with some modifications in the 
interval led to the conviction that under the conditions the 
cotyledons are practically exhausted in somewhat less than 10 
days. It might be pointed out that the removal of the coty- 



\nn. Mo. Bot. Gard.. Vol. 7, 1920 



Plate 7 



Safes** * 





' - ^ *< 



DUGGAR— FOOD RESERVE IN COTYLEDONS 

(Canada field peas, see tab e 1) 



1920] 



DUGGAR — VALUE OF FOOD RESERVE IK COTYLEDONS 



293 



TABLE I 

THE EFFECTS OF THE RESERVE FOOD SUPPLY ON THE GROWTH OF 

SEEDLINGS 



Cult, 
no. 



1 

2 

3 
4 
5 

6 



Field corn, 10 plants, 
average of duplicate cultures 



Time of 
excision of 
endosperm 



After 2 days 
After 5 days 
After 7 days 
After 8 days 
After 9 days 
After 10 days 



Canada field peas, 10 plants, 
average of duplicate cultures 



Total 




green wt., 


Cult. 


grams 


no. 



32.72 

30.18 
34.02 
30.11 
36.11 
36.98 



1 
2 
3 
4 
5 
6 



Time of 
excision of 

cotyledons 



Aft er 2 days 
Aft er 3 days 
Aft er 4 days 
After 5 days 
After 7 days 
Control, uncut 



Total 

green wt., 

grams 



5.75 
10.12 

17.70 
19.82 
24.30 
25.51 



ledons was done in all cases with the greatest care, so that no 

injury to the seedling would result. The excision was made 

at a point beyond the stalk of the cotyledons. In the case of 

corn the results are a little irregular. Nevertheless, there is 

the suggestion that the removal of the main carbohydrate food 

supply is not so important a factor in depressing the growth of 

the young plant. From subsequent incidental experiments I 

am convinced that there is some effect, but i b is neither so marked 

as in the case of the peas nor does it seem to be so permanent, 

that is, the effect is not so striking during the further develop- 
ment of the plant. 

During the summer of 1919 1 an attempt was made to sub- 
stitute for the loss of the cotyledons in the; case of the peas by 
the addition of certain organic nitrogen-containing nutrients, 
and especially by the addition of glycocoll, alanin, sodium 
asparaginate, and sodium nucleinate. These experiments were 
carried out under the most favorable conditions for the growth 

1 This series of experiments and other supplementary studies not yet concluded 
were carried out at the Coastal Laboratory of the Carnegie Institution of Washing- 
ton, Carmel, California, and the writer takes this occasion to acknowledge his in- 
debtedness to the Director of Botanical Research, Doctor D. T. MacDougal, for 
placing at his disposal the facilities of the laboratory and for his cordial cooperation. 



(Vol. 7 



294 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Qrttn Hi. in Grams 




gotyUulons gKCtsccl 



Stnmtliuttlu %da.%. 7cCus. Coxtnl 



Fig. 1. Green weight quantities of Canada field peas 
in various solutions as affected by cotyledon excision. 



of peas, at a mean temperature of about 15.6° C. The technique 
was the same as above described, but the use of organic sub- 
stances in the solutions made it desirable to renew the solutions 
every 3 or 4 days in order to reduce or control bacterial action. 
The complete results are shown in table n, and in fig. 1. It 
will ultimately be necessary to extend the use of organic sub- 
stances and to repeat this work under pure culture conditions. 
In fact, a small series of experiments in this direction has already 
been performed, but inasmuch as these are to constitute a part 
of a more extended study I will confine myself here to a brief 
discussion of the data presented below. 

In general it will be seen that whatever the medium employed 
as a nutrient solution the removal of the cotyledons is shown 
by marked depression in the growth rate. The mineral nutrient 
solution employed in this case is that which I have in a previous 



1920] 



DUGGAR — VALUE OF FOOD RESERVE IN COTYLEDONS 



295 



TABLE II 

THE EFFECTS OF THE EXCISION OF THE COTYLEDONS UPON THE GROWTH 

OF CANADA FIELD PEAS 



Cult. 


Culture 
medium 


Time of 
excision 


Total 
gr. wt. 


Gr. wt. 

of tops 


(Jr. wt. 

of roots 


Per cent 
rel. to 

cor. cult., 
sol. C* 


Per cent 
rel. to 


no. 


10 plants, weight in grams 


cult. 4 


i 

2 
3 
4 


I 

Solution C 


Immediately 
After 2 days 
After 7 days 
Control, uncut 


10.26 
14.90 
17.50 
22.11 


4.97 

8.75 

11.64 

14.40 


5.29 
6.15 
5.86 
7.71 


100 
100 
100 
100 


46 

67 

79 

100 


5 
6 

7 
8 


Solution C 

+ 
glycocoll 

(M/100) 


Immediately 
After 2 days 
After 7 days 
Control, uncut 


12.28 
16.90 
19.81 
19.29 


7.38 
10.40 
13.93 
13.45 


4.90 

6.50 

5.88 
5.84 


120 
113 
113 

87 


56 
76 

90 

87 


9 

10 
11 
12 


Solution C 

+ sodium 

asparaginate 

(M/100) 


Immediately 
After 2 days 
After 7 days 
Control, uncut 


6.57 
11.04 
16.44 
17.26 


3.92 

6.68 

10.09 

11.52 


2.65 
4.36 
6.35 

5.74 


64 
74 

94 

78 


30 
50 
74 

78 


13 
14 
15 
16 


Solution C 

alanin 
(M/100) 


Immediately 
After 2 days 

After 7 days 
Control, uncut 


7.53 
10.69 
16.56 
17.86 


4.62 
6.92 

10.45 
12.00 


2.91 

3.77 
6.11 

5.86 


73 

72 

95 
81 


34 
48 
75 
81 


17 

18 

19 
20 


Solution C 

+ sodium 

nucleinate 

( Xo per cent) 


Immediately 
After 2 days 
After 7 days 
Control, uncut 


14.38 
18.32 

22.85 

25.43 


7.29 
10.13 
14.57 
15.09 


7.09 

8.19 

8.28 

10.34 


140 
123 
131 
115 


65 

83 

103 

115 



* Per cent relative to corresponding culture, with respect to the time of excision 
of cotyledons, in the unmodified solution C. 

paper (Duggar, '20) called solution C, which is one of the "best" 
combinations developed by Livingston and Tottingham ('18), 
and this solution has been shown to contain a favorable concen- 
tration of potassium nitrate for the growth of wheat and peas. 
Nevertheless, in spite of the fact that at the time of the excision 
of the cotyledons (even in those excised after only 2 days) there 
was considerable green leaf tissue in the seedlings, still the 
growth was weak, and at the end of 24 days the plants weighed 
less than one-half the control. The addition of glycocoll and 
of sodium nucleinate increased the growth quantities materially 
in the corresponding cultures, while the addition of sodium 
asparaginate or alanin was slightly depressing. The depressing 



|Vol. 7 



296 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



action, however, may have been due to a small amount of bac- 
terial decomposition products. Even in the presence of sodium 
nucleinate, as in culture 17, the amount of growth when the 
cotyledons were excised after 2 days is much less than in the 
control (cotyledons uncut) in solution C, culture 4. The data 
seem to indicate that no proper nutrient substitute for the coty- 
ledons has been found in these organic substances. Some addi- 
tional experiments in which sugar was used in connection with 
nitrogen-containing substances have not served to change ma- 
terially the conclusions which may be drawn. In other experi- 
ments urea and nucleic acid were used, but neither of these 
has been as favorable as sodium nucleinate or glycocoll. It is 
true, however, that sodium nucleinate has increased more than 
any other compound thus far used the growth quantities in the 
cultures lacking cotyledons. 

The importance of the cotyledons in the vigorous development 
of the seedling is an ancient observation. Bonnet (1754) demon- 
strated that beans and buckwheat grew less rapidly when the 
cotyledons were cut off, and more important still, he observed 
the persisting effect of this early difference, stating the matter 
in the following words: "La meme difference, ou une difference 
analogue, a subsiste entre ces Plantes pendant toute la duree 
de l'accroissement. II a toujours ete facile de distinguer les 



des 



•>■> 



Sachs ('59) observed the same fact a century later while 
devoting more attention to the physiology of absorption and 
nutrition. Discussing numerous experiments designed to deter- 
mine the interdependence of organs and tissues in the embryo, 
Van Tieghem (73) refers incidentally to the problem here dis- 
cussed. 

While Schmid ('94), Hannig ('04), and Smith ('07) have con- 
tributed many interesting observations regarding the nutrition 
of the embryo and the capacity of different parts to develop or 
regenerate, these facts do not closely relate to the present in- 
vestigation. Dubard and Urbain ('13), however, emphasize the 
favorable effects of the endosperm of certain grains in the early 
stages of germination. They directed their work primarily 
toward determining the capacity of the embryos to develop in 
the absence of the endosperm. 



1920 



DUGGAR — VALUE OF FOOD RESERVE IN COTYLEDONS 297 



Recently Andronescu ('19) has sought to determine the im- 
portance of the endosperm and scutella in Zea Mays and at 
the same time he has endeavored to find a substitute for these, 
also to follow in heredity any effects observed. While demon- 
strating that normal plants develop without endosperm, his 
use of the term normal is a relative one, and. he concludes with 
the statement, "We cannot deny, however, that the presence of 
endosperms is beneficial in the process of germination, as well 
as in the further development of the plants." 

The writer proposes to continue these investigations with 
plants grown under sterile conditions in the hope of determining 
more definitely the nature of the special nutrient or growth- 
inducing substance furnished by the cotyledons. At present 
one of several explanations for the failure to substitute readily 
for the cotyledons may be given: (1) it is conceivable that a 
combination of organic nutrients including several amino acids 
may be necessary for normal growth; (2) that penetration of 
organic substances may be slow and difficult; and (3) that the 
cotyledons may contain a vitamine requisite for the vigorous 
development of the plant. In any case the results so far ob- 
tained, as well as the observations of other investigators, indi- 
cate that the depression of growth accompanying the excision 
of the cotyledons is marked in the case of peas and other plants 
with fleshy seed-leaves, and that the influence of excision extends 
throughout the growth period of the plants. 1 



Bibliography 

Andronescu, D. I. ('19). Germination and further development of the embryo 
of Zea Mays separated from the endosperm. Am. Jour. Bot. 6: 443-452. 
pt. 41. 1919. 

Bonnet, C. (1754). Recherches sur l'lisage des feuilles dans les plantes. 1754. 
[See pp. 236-242.] 

Dubard, M., et Urbain, J. A. ('13). De l'influence de l'a butnen sur le developpe- 
ment de l'embryon. Compt. Rend. Acad. Paris 156: 1086-1089. 1913. 

1 Since this paper was written there has appeared another article of considerable 
interest dealing with the effect of the endosperm upon the growth of the embryo 
(Urbain, A., Influence des matieres de reserve de r albumen de la graine sur le devel- 
oppement de Tembryon. Rev. Gen. Bot. 32: 125-139, 165-191. U fig. 1920.). 
In addition to careful observations on the effects of the excision of the endosperm 
on the development of a number of plants, a careful comparative study was made of 
internal structure. 



(Vol. 7 



298 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



Duggar, B. M. C19). Nutritive value of food reserve in cotyledons. Carnegie 
Inst. Wash., Year Book (Ann. Rept. Dir. Dept. Bot. Res.) 18: 81-82. 1919. 

, C20). Hydrogen ion concentration and the composition of nutrient 



solutions in relation to the growth of seed plants. Ann. Mo. Bot. Gard. 7: 
1-49. /. 1-7. 1920. 

Hannig, E. ( , 04). Zur Physiologie pflanzlicher Embryonen. Bot. Zeit. 62: 45-80. 
pi 3. 1904. 

Livingston, B. E., and Tottingham, W. E. ('18). A new three-salt nutrient solu- 
tion for plant cultures. Am. Jour. Bot. 5: 337-346. 1918. 

Sachs, J. ('59). Physiologische Untersuchung uber die Keimung der Schmink- 
bohne (Phaseolus multiflorus). K. Akad. Wiss., matb.-naturw. CI., Sitz- 
ungsber. 37: 57-119. pi 1-3. 1859. 

Schmid, B. ( , 94). Ueber die Lage des Phanerogamen-Embryo. Bot. Centralbl. 58: 
1-7. 1894. 

Smith, Louie H. C07). Beobachtungen uber Regeneration und Wachstura an 
isolierten Teilen von Pflanzenembryonen. Inaug. Dissertation. 86 pp. 4 pis. 
Halle, 1907. 

Van Tieghem, P. (73). Recherches physiologiques sur la germination. Ann. 
Sci. Nat. Bot. V. 17: 205-224. 1873. 



TITRATION CURVES OF CERTAIN LIQUID CULTURE 

MEDIA 

JOANNE L. KARRER 

Research Assistant to the Missouri Botanical Garden 



AND ROBERT W. WEBB 



Lackland 



Washington University 



A study of the growth of various fungi and of the germina- 
tion of various fungous spores with reference to H-ion concen- 
tration has involved a determination of the titration curves of 
several culture media. It is the purpose of this paper to present 
briefly the data obtained with these nutrient solutions, which 
have been the basis of subsequent work in this laboratory. 

The formulae for the media employed are as follows: 

Beet decoction. — Prepared according to the method outlined 
by Duggar, Severy, and Schmitz ('17). This consists essen- 
tially of 370.4 gms. of sugar beets per liter of distilled water, 
autoclaved at 15 pounds for one hour, and then filtered. 

Czapek's solution.— MgS0 4 +7 H 2 0, .5 gm.; KH 2 P0 4 , 1.0 gm.; 
KC1, .5 gm.; NaN0 3 , 2.0 gms.; FeS0 4 , .01 gm.; cane sugar, 
30.0 gms.; distilled H,0, 1000 cc. (Zeller, Schmitz, and Duggar, 
'19). 

Peptone solution. — Twenty gms. bacto-peptone in 1000 cc. 
H 2 0. 

Pfeffer's solution.— KH 2 P0 4 , 5.0 gms.; MgS0 4 +7 H 2 0, 2.5 
gms.; NH 4 N0 3 , 10.0 gms.; FeS0 4 , trace; cane sugar, 50.0 gms.; 
distilled H 2 ; 1000 cc. (Pfeffer, '95). 

Richards' solution. — KH 2 P0 4 , .5 gm.; KN0 3 , 4.0 gms.; MgS0 4 
+7 H 2 0, .5 gm.; NH 4 N0 3 , 10.0 gms.; FeS0 4 , trace; cane sugar, 
30.0 gms.; distilled H 2 0, 1000 cc. (Richards, '97). This formula 
differs from that of the original by a reduction in the amount 
of MgS0 4 . The amount of precipitate produced in the more 
alkaline solutions was found to depend largely upon the amount 
of magnesium present, and, since it is important to keep the 
amounts of the constituents in the solutions as nearly equal as 
possible while varying the H-ion concentration, it seemed desir- 
able to reduce the amount of MgS0 4 to .5 gm. per liter. Such 



Ann. Mo. Bot. Gard., Vol. 7, 1920 



(299) 



[Vol. 7 



300 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



a reduction has been found by Duggar (unpublished data) to 
have no appreciable effect upon the growth of fungi. 

The nutrient solutions were all prepared by adding 600 cc, 
instead of 1000 cc, of distilled water. This method allowed 
dilutions of the various solutions by additions of regulated 
amounts of acid or alkali and water for the adjustment of vari- 
ous H-ion concentrations without affecting materially the con- 
centrations of the nutrient salts or constituents. 

It is possible to obtain a wide range in the H-ion concentra- 
tions of the solutions by adding an alkali and a mineral acid 
in successively increasing quantities. The addition of the mono-, 
the di-, or the tri-basic potassium phosphate as suggested by 
Zeller, Schmitz, and Duggar ('19) was found to give results 
within a certain range, but additions in too large amounts were 
necessary to produce extreme concentrations. Therefore, since 
such a decided variation in the composition of the nutrient 
solutions undoubtedly would have existed, it seemed undesir- 
able to adopt this method in the experiments under investi- 
gation. 

From preliminary experimentation relative to the adjust- 
ment of H-ion concentration in media of alkaline reactions, 
the most satisfactory results were obtained by using N /5 KOH, 
and an alkali of this strength has been employed in all of the 
experiments except in the case of Czapek's solution. Here, 
the buffer action was less than that in the other nutrient solu- 
tions, consequently N/20 concentration was more conveniently 
and accurately applicable. In all the experiments, however, 
N/5 HC1 was favorable for varying the reactions on the acid 
side. In the case of Czapek's solution, the reactions were also 
varied by means of N/1 H 3 P0 4 and N/20 NaOH. These re- 
sults so nearly paralleled those obtained with the above acid 
and alkali that it was deemed unnecessary to continue this 
aspect of the experiment with the other nutrient solutions. 

Inasmuch as it is generally admitted that sugars readily 
react with acid or alkali when heated under pressure, the nutri- 
ent solutions and the acid and the alkali were sterilized sep- 
arately. Thirty cc. of the nutrient solution together with the 
desired amount of water, as indicated in the tables, were put 
into small flasks, plugged, and sterilized. After cooling, the 



1920] 



KARRER AND WEBB — TITRATION CURVES OF CULTURE MEDIA 301 







(Vol. 7 
302 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



cultures were removed to a culture chamber, and definite amounts 
of sterile acid and alkali were added by means of sterile gradu- 
ated pipettes. The final volume of each culture was 50 cc. 
and represented a dilution of the constituents comparable with 
that in the original nutrient solutions. The solutions were 
allowed to stand for 24 hours in order to reach a state of equili- 
brium, and, at the expiration of this period, H-ion determinations 
were made according to the colorimetric method of Clark and 
Lubs ('17), all determinations being made at room temperature. 
(See figs. 1-5 and tables i-ii). 

Due to the presence of color in the beet decoction and in the 
peptone solution, it was necessary to use a colorimeter for the 
H-ion determinations. A Duboscq type was used in this par- 
ticular work, the detailed method of which has been described 
by Duggar ('19). 

In all of the mineral salt solutions, a certain amount of pre- 
cipitate occurred upon the addition of alkali, the amount in- 
creasing with increase of added alkali. No such phenomenon 
was evidenced in the alkaline cultures of the beet decoction 
or of the peptone solution. The precipitation referred to com- 
menced with culture No. 29 in Pfeffer's solution, No. 27 in Rich- 
ards' solution, and very faintly in No. 23 of Czapek's solution. 
On the other hand, there was no precipitation whatever in the 
acid cultures of any of the various media. 

As the reaction of the beet decoction passed from acid to 
alkaline, there was noted a decided color change from pale 
yellow to amber, and a slight cloudiness was perceptible begin- 
ning with culture No. 42. 

The titration curve obtained with 2 per cent bacto-peptone 
agrees closely with the curves obtained by Clark and Lubs ('17) 
with Witte peptone, falling, as one would expect, between I he 
curves representing concentrations of 1 and 5 per cent. 

The initial P H of the original culture solutions varied, varia- 
tions of several tenths not being infrequent despite the most 
careful technique during preparation and the use of highest 
purity chemicals. Since the highest purity mono-basic potas- 
sium phosphate obtainable had a very high acidity due to the 
impurities present, the salt was recrystallized until the Sorensen 
coefficient of P H 4.529 for a 1/15 molecular solution was ob- 
tained. 



1920] 



KARRER AND WEBB — TITRATION CURVES OP CULTURE MEDIA 



303 



No. 



1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 
14 
15 
16 
17 
18 
19 
20 
21 

22 

23 
24 
25 
26 

27 

28 

29 
30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

40 

41 

42 

43 

44 

45 

46 

47 
48 
49 



TABLE I 

TITRATION DATA FOR VARIOUS LIQUID MEDIA 



Solution 
(cc.) 



30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 



N/5 
HC1 

(cc.) 



N/5 



20.00 

15.00 

10.00 

9.50 

9.00 

8.50 

8.00 

7.50 

7.00 

6.50 

6.00 

5.50 

5.00 

4.50 

4.00 

3.50 

3.00 

2.50 

2.00 

1.50 

1.00 

.75 

.50 

.25 

0.00 



(cc.) 



.25 

.50 

.75 

1.00 

1.50 

2.00 

2.50 

3.00 

3.50 

4.00 

4.50 

5.00 

5.50 

6.00 

6.50 

7.00 

7.50 

8.00 

8.50 

9.00 

9.50 

10.00 

15.00 

20.00 



Dist 



KOH I H,0 



(cc.) 



00.00 
05.00 
10.00 
10.50 
11.00 
11.50 
12.00 
12.50 



13.50 
14.00 
14.50 
15.00 
15.50 
16.00 
16.50 
17.00 
17.50 
18.00 
18.50 
19.00 
19.25 
19.50 
19.75 
20.00 
19.75 
19.50 
19.25 
19.00 
18.50 
18.00 
17.50 
17.00 
16.50 
16.00 
15.50 
15.00 
14.50 
14.00 
13.50 
13.00 
12.50 
12.00 
11.50 
11.00 
10.50 
10.00 
5.00 
0.00 



Total 
(cc.) 



50 
50 
50 
50 
50 
50 
50 
50 



13.00 50 



50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 



Hydrogen ion concentration, Ph 



Beet 
decoc- 
tion 



1.2 

1.2 

1.6 



1.7 



19 



2.1 



2.3 



2.6 



3.1 



3.4 
3.6 

3.8 



4.4 
4.5 
4.8 
5.0 
5.2 
5.4 
5.6 
6.4 
7.0 



8.8 



9.8 



10 0+ 



2.0 
2.0 



2.2 



2.4 



2.8 



4.6 



7.0 

7.5 

7.7 
7.9 
8.1 
8.2 
8.2 
8.3 
8.3 
8.4 
8.5 
8.6 
8.6 
8.7 
8.7 
8.7 
8.7 
8.7 
8.8 
8.9 
9.1 



4.4 



4.9 



5.4 



6.4 



7.0 



7.4 



7.8 



8.2 



8.8 



9.2 



9.6 



10.0 



10.0 



10.0+ 



10.0+ 



10.0+ 



Rich- 


Pep- 


Pfef- 


ards 


tone 


fer 


1.2 


1.4 


1.4 


1.2 


2.0 


1.5 


1.4 


3.0 


1.5 


1.4 




1.6 


1.5 


3.2 


1.7 


1.5 




1.7 


1.5 


3.4 


1.8 


1.6 




1.8 


1.6 


3.6 


1.9 


1.6 




1.9 


1.7 


3.8 


1.9 


1.7 




2.0 


1.7 


4.0 


2.1 


1.8 




2.1 


1.8 


4.2 


2.1 



2.3 
2.3 
2.3 
24 

2.7 
2.8 



3.2 



4.3 



5.3 



5.6 
6.2 
63 
6.4 
6.6 
6.6 
6.8 
69 
7.0 

7.1 

7.2 

7.3 
7.4 

7 6 

7.8 
80 
8.2 

8.3 

8.4 

9.0 
9.6 



304 



[Vol. 7 



ANNALS OP THE MISSOURI BOTANICAL GARDEN 



TABLE II 

TITRATION DATA FOR CZAPEK'S SOLUTION 



No 



1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 
14 
15 
16 
17 
18 
19 
20 
21 
22 
23 
24 

25 
26 
27 
28 
29 
30 
31 
32 

33 
34 
35 
36 
37 
38 
39 

40 
41 
42 
43 
44 
45 



Czapek 
sol. 
(cc.) 



30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 

30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
30 



Acid 
(cc.) 



20.00 
15.00 
10.00 
9.50 
9.00 
8.50 
8.00 
7.50 
7.00 
6.50 

6.00 
5.50 
5.00 
4.50 
4.00 
3.50 
3.00 
2.50 
2.00 
1 . 50 
1.00 
.50 
00 



Alkali 
(cc.) 






.50 
1.00 
1.50 

2.00 
2.50 
3.00 
3 . 50 
4.00 
4.50 
5.00 
5.50 
6.00 
6.50 
7.00 
7.50 
8.00 

8 . 50 
9.00 

9 . 50 
10.00 
15.00 
20.00 



Dist. 
H 2 
(cc.) 



0.00 

5.00 

10.00 

10.50 

11.00 

1 1 . 50 

12 . 00 

12.50 
13.00 

13 . 50 
14.00 

14 . 50 

1 5 . 00 
15 . 50 

16 . 00 

16. 50 
17.00 
17.50 
18.00 
18.50 
19.00 
19.50 
20.00 

19.50 
19.00 
1 8 . 50 
18.00 
17.50 
17.00 
16.50 
16.00 
15 . 50 
15.00 
14.50 
14.00 
13.50 
13.00 
1 2 . 50 
12.00 
1 1 . 50 
11.00 
1 50 
10.00 
5.00 
0.00 



Total 
cc. 



50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 

50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 
50 

50 
50 
50 



H-ion concentration, Ph 



N/5 HC1 
N/20 KOH 






1.2 
1.3 

1.4 
1.4 

1.5 
1.5 
1.5 
1.6 

1.6 

1.6 
1.7 
1.7 
1.8 
1.9 
1.9 
2.0 
2.1 
2.2 
2.3 
2.4 
2.5 
2.9 
4.7 
5.7 
6.0 
6.2 
6.4 
6.6 

6.7 
6.8 
6.9 
7.0 
7.1 
7.2 
7.4 
7.5 
7.6 
7.7 

8.0 

S.6 
8.9 
9.1 
9.2 
9.8 
9.8 



N/1 H 3 PO, 
N/20 NaOH 



1.6 
1.6 
1.6 

1.7 
1.7 
1.7 
1.7 

1.8 
1.8 
1.8 
1.9 

1.9 

2 
2 
2.1 
2.2 
2.3 
2.3 
2 4 

2.6 

4.4 
5.0 

5.9 
6.2 
6.3 
6.5 
6.6 
6.8 
6.8 
7.0 
7.1 
7.2 
7.4 
7.6 
7.7 
7.9 
8.2 
8.6 
8.7 
9.0 
9.2 



1920 J 



KARRER AND WEBB — TITRATION CURVES OF CULTURE MEDIA 305 



The writers take pleasure in extending thanks to Dr. B. M. 
Duggar for many suggestions and criticisms ; and to Dr. George T. 
Moore for the privileges and facilities of the Missouri Botanical 
Garden. 

Graduate Laboratory, Missouri Botanical Garden. 



Literature 

Clark, W. M., and Lubs, H. A. ('17). The coloriraetric determination of hydrogen 
ion concentration and its applications in bacteriology. I. Jour. Bact. 2: 
1-34. /. 1-4. 1917. 

Duggar, B. M. ('19). The micro-colorimeter in the indicator method of hydrogen 
ion determination. Ann. Mo. Bot. Gard. 6: 179-381. 1919. 

Duggar, B. M., Severy, J. W., and Schmitz, H. ('17). Studies in the physiology 

of the fungi. IV. The growth of certain fungi in plant decoctions. Ibid. 
4: 165-173. /. /->$. 1917. 

Pfeffer, W. ('95). Ueber election organische Nahrstoffe. Jahrb. f. wiss. Bot. 28: 
205^268. 1895. 

Richards, H. M. ('97). Die Beeinflussung des Wachstmus einiger Pilze durch 
chemische Reize. Ibid. 30: 665-688. 1897. 

Sorensen, S. P. L. ('09). Enzymstudien II. Ueber die Messung und die Bedeutung 
der Wasserstoffionenkonzentration bei enzymatis 3hen Prozessen. Biochem. 
Zeitschr. 21: 129-304. 1909. 

Zeller, S. M., Schmitz, H., and Duggar, B. M. C19). Studies in the physiology 
of the fungi. VI. Growth of wood-destroying fungi in liquid media. Ann. 
Mo. Bot. Gard. 6: 137-142. 1919. 



THE USE OF "INSOLUBLE" SALTS IN BALANCED 

SOLUTIONS FOR SEED PLANTS 



B. M. DUGGAR 

Physiologist to the Missouri Botanical Garden, in Change of Graduate Laboratory, 

Professor of Plant Physiology in the Henry Shaw School of Botany of 

Washington University 

In this paper it is proposed to give the results of several series 
of experiments designed primarily to determine the possible 
value of certain relatively insoluble salts in furnishing the nec- 
essary ions for the growth of seed plants. By means of such 
salts it will also be attempted to secure favorable combinations 
of the necessary ions. Throughout this discussion "insoluble" 
may be used in a very general sense, to include many salts soluble 
only to a comparatively slight degree, or with difficulty soluble, 
in water at from 15 to 25° C. 

It is well known that in the soil a relatively small part of 
the salts ordinarily designated mineral nutrients is present in 
soluble form. There is, in general, a very considerable reserve 
or "unavailable" supply of the less readily soluble salts of such 
elements as K, Ca, Mg, Fe, S, and P. Nitrates are generally 
present only in low concentration and the surplus nitrogen 
supply is usually in the form of organic compounds. It has 
seemed to the writer eminently desirable to determine, there- 
fore, if a favorable nutrient solution for seed plants may not 
be arranged from combinations of some of these insoluble salts, 

thus in some measure imitating the chemical relations in the 
soil. 

In favor of this endeavor it might be argued that should this 
prove possible it would only be necessary to add to the culture 
vessel a surplus of the substances required. A small amount of 
that added would go into solution immediately and when an 
equilibrium were attained the absorption oi any ions by the root 
would be compensated for by further solution of the substances 
furnishing these ions, and thus the equilibrium might be main- 
tained and the concentration kept fairly constant over con- 
siderable intervals. Obviously, it would be impracticable to 
furnish nitrate as an insoluble compound, since the nitrates of 

ANN. Mo. BOT. G.UtD , VOL. 7, 1920 (307) 



[Vol. 7 



308 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



the bases required are all soluble to a high degree. If, there- 
fore, nitrogen is furnished as N0 3 , the salt furnishing this ion 
would necessarily be added periodically, and to this extent the 
concentration would change from day to day. Relatively in- 
soluble salts of ammonium are obtainable, however, such as 
MgNH 4 P0 4 , and this salt has been employed in some of the 

experiments. 

In the various experiments which have thus far been under- 
taken the sources of Ca are as follows: CaS0 4 +2H 2 0, CaCO,, 
Ca,(PO.)„ and CaHP0 4 +2H 2 0; of Mg: MgS0 4 +7H 2 0, MgCO,, 
Mg,(PO,),+8H,0, and MgNH 4 P0 4 +6H 2 0; of K: KNO, and 
K 3 P0 4 ; of Fe: FeP0 4 +4H 2 0, FeC 2 4 +2H 2 0, ferric citrate, 
and "soluble ferric phosphate." Certain other salts which 
might have been employed to advantage were not available at 

the time. 
Among the more soluble of the salts included in this category 

are CaS0 4 +2H 2 having a solubility of 0.241 and 0.222 in 100 
parts of water at 0° and at 100° C. respectively, and among the 
more insoluble salts are FeP0 4 +4H 2 and CaC0 3 . One of the 
chief reasons for burdening the experiments with such a variety 
of substances may be found in the fact of their diverse solu- 
bilities; and inasmuch as the antagonistic relations of the ions 
in respect to the plant require consideration and are involved 
with osmotic and nutritive relations, such a variety of combina- 
tions was necessary in order to feel assured that some of the 
results obtained might be among the most favorable that such 
types of combinations could yield. 

It is obvious that at no instant is the exact concentration of 
any salt known in such combinations, except approximately, 
in cases where soluble salts were added. However, the total 
concentration is readily determinable, likewise the electrolytic 
conductivity of the solution. One may also estimate the partial 
concentrations. When soluble salts are employed in nutrient 
solutions, the proportion of the ions is disturbed from the mo- 
ment that the roots are introduced, and there is a progressive 
decrease in concentration until the solution is renewed. Like- 
wise the differences in the proportion of the ions, determined, 
of course, by the differential absorption rates, are not continually 
reestablished by any form of "reserve." It would, of course, 



1920] 



DUGGAR — INSOLUBLE SALTS IN BALANCED SOLUTIONS 309 



be possible to effect a circulation of fresh nutrient solutions 
where soluble salts are employed, but any operation of this 
nature would be impracticable in most of our experimental 
work. After all, the problem is to obtain a combination of 
salts favorable to a high degree which may be employed in 
practically any type of experiment where the desire is to main- 
tain the plant under satisfactory physiological conditions. If 
the labor involved is great in the one case and small in the other 
the one involving the less labor will, of course, be selected as 

the more practicable. 

Before describing in detail the methods and experiments to 
be discussed it should be pointed out that the nutrient solution 
developed by Crone ('03) and considered by him to have certain 
advantages over the Pfeffer solution, contained, in addition to 
KN0 3 , MgS0 4 , and CaS0 4 , ferrous phosphate and tribasic cal- 
cium phosphate. The two salts last mentioned are, of course, 
relatively insoluble and were used by Crone with the idea of 
diminishing the chlorosis which he attributed to the excess of 
soluble phosphate and the low content of iron. Later, how- 
ever, Benecke ('09) was unable to substantiate the claims made 
by Crone as to the benefits to be derived from the type of solu- 
tion which the latter had formulated. 

The experiments on which the data in table 1 are based were 
carried out in the experimental greenhouse at the Missouri Bo- 
tanical Garden during April, 1920. The msthods employed were 
in fairly close accord with those previously described. Glass 
tumblers of 250 cc. capacity were used with 240 cc. of nutrient 
solution. To these containers were fitted eorks 7.5 cm. in diam- 
eter and 0.7 cm. in thickness, arranged with holes for the inser- 
tion of the roots of the seedlings, and with an extra hole to facili- 
tate the addition of water lost by transpiration. The seedlings 
were germinated over water, and in order to insure the greatest 
possible uniformity in size a selection of these was made when 
the shoots were about 2 cm. in height. Each cork was held in 
position by a stout rubber band passing around the tumbler 
lengthwise. The tumblers were covered as usual to protect the 
roots from the light, and the cultures were then freely and equally 
exposed on a lattice bench in the greenhouse. The variety of 
wheat employed throughout was the Pacific Coast Blue Stem, 



[Vol. 7 



310 ANNALS OF THE MISSOURI BOTANICAL GARDEN 



supplied by the Plant Introduction Garden of the Bureau of 
Plant Industry, Chico, California. The corn was a standard 
field strain of yellow Dent. 

The results shown in tables n and in 1 were obtained at Car- 
mel, California, during July and August of the same year. At 
Carmel the cultures were arranged on lattice tables in the open. 
The average temperature was about 15.6° C. and the average 
daily evaporation from a standard spherical porous cup at- 
mometer about 15 cc. Table n represents cultures prepared 
exactly as in table i except that glass beakers, of the same ca- 
pacity, were used instead of tumblers. The data in table in 
are from experiments closely paralleling those represented by 
table ii except that in the former the containers used were one- 
quart preserving jars (Economy style). This type of jar proved 
most convenient in this work, since the mouth of the jar is large, 
taking the same corks as used in the tumblers and beakers. 
Moreover, the spring clips which accompany these jars afford 
a handy method of fastening the cork to the jar so that the 
seedlings are not readily disturbed. The use of the larger con- 
tainers in this case explains the larger quantities of salts or 
solutions employed, and, of course, vessels of this capacity per- 
mit the experiments to be continued over a longer period of 

time. 
Inasmuch as certain cultures in each of these series contained 

not only a full mineral "nutrient" solution but also some citrate, 

it seemed well to arrange all the solutions and then let them 

stand two days in case some evidence of fermentation might 

develop. This occurred in certain cases especially in the second 

and third series, but afterwards cleared up. The significance of 

this will be discussed in a later paper in which the physical 

characteristics of nutrient solutions in general will receive special 

consideration. 

In all cases where readily soluble salts were used in these 
experiments the initial quantities in the different series varied 



considerably 



added from time to time, 



1 This work was done at the Coastal Laboratory of the Carnegie Institution of 
Washington. The writer is pleased to make acknowledgment of the facilities and 
cooperation extended by Doctor D. T. MacDougal, Director of Botanical Research, 
and of the courtesies of other members of the staff. 



1920] 



DUGGAR — INSOLUBLE SALTS IN BALANCED SOLUTIONS 311 



and these facts are brought out in the special explanations 
given in connection with the particular tables. 

In table i there are given in the second and third columns 



(under "concentration," i and n) the quantities of the salts 
used, these being expressed in grams of the pure salt or in cc. 
of a standard stock solution. The concentration numbers occur 
again in the fifth column, indicating to which concentration 
the data in the remaining columns refer. When given in grams, 
the quantities indicated were used in 240 cc. of water, and no 
change of these constituents was made throughout the interval 
of growth. 

The quantities given in cc. also require explanation. KN0 3 : 
the stock solution employed for every culture in which this 
salt occurs except No. 15 contains 35 grams KN0 3 in 1000 cc. 
of water, and the use of 10 cc. per culture of 240 cc. gives a 
concentration of this salt in the culture solution approximately 
three times as great as in solution B (No. 15 in this table). It 
is approximately two-thirds as strong as the concentration of 
KN0 3 in one of the "best" cultures of Livingston and Totting- 
ham ('18), that is, Red, referred to in my earlier paper (Duggar, 
'20) as solution C. Moreover, in the two cultures (No. 12 and 
No. 14) in which Mg(N0 3 ) 2 or NaN0 3 was substituted for KNO 
the strength of the solution was such as to afford a quantity 
of N0 3 equivalent to that of the KN0 3 in all cultures except 
No. 15. MgS0 4 : the stock solution, 12 grams in 1000 cc. of 
water, is the same as that used in solution B (No. 15 of this 
series). The concentration of soluble ferric phosphate is like- 
wise made the same as in solution B (No. 15 of this table). The 
control solution in this series is solution B, No. 15 of the table, 
previously described in detail, as noted above. Additions of 
10 cc. of KN0 3 were made to each culture (240 cc.) containing 
this salt at intervals of 7 days, and at the same time the solu- 
tion in No. 15 (solution B, control) was renewed. 

A glance at table 1, and more especially a study of fig. 1 (wheat), 
indicates that the differences between the two "concentrations" 
or strengths of solutions are within the probable limits of varia- 
tion commonly found in duplicate cultures. The average of 
the two similar control cultures in solution B (No. 15) is ex- 
ceeded by No. 2. The latter culture differs from the control 



^ 



|Vol. 7 



312 



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DUGGAll — INSOLUBLE SALTS IN BALANCED SOLUTIONS 



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ANNALS OF THE MISSOURI BOTANICAL GARDEN 



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1920] 



DUGGAR — INSOLUBLE SALTS IN BALANCED SOLUTIONS 



315 



I Green weight 

In grams 
to 
o 



Wheat 



o 




Fig. 1. Yield of wheat in solutions of relatively insoluble salts. Continuous 
line is concentration I and broken line concentration n (see table i). 



(1) in the excess of CaSO< (solid phase present), (2) in the sub- 
stitution of the "insoluble" tribasic magnesium phosphate for 
the soluble MgS0 4 , and (3) in the greater content of KN0 3 . It 



will be noticed also that cultures 4, 8, and 10 approach the 



value of the control, and all of these contain the soluble ferric 
phosphate, combined with various calcium and magnesium com- 
pounds. No culture containing either KjPO* or iron oxalate has 
yielded satisfactorily. Similarly, MgCO a in the combinations 
employed would seem to be less depressing than K 3 P0 4 but still 
unfavorable. Among the cultures mentioned as giving the higher 
yields no striking peculiarity was noted except in the case of 
No. 10, in which there was pronounced tillering at a relatively 
early period. 

With corn many cultures are ahead of the control, No. 15, 
and those in advance are again generally the cultures contain- 
ing soluble ferric phosphate, though the differences between 
the pairs containing FePO* and the salt of :ron just mentioned 



316 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 




Fig. 2. Yield of corn in solutions of relatively insoluble salts. Continuous 
line is concentration i and broken line concentration h (see table i). 



are not so striking as with wheat. Nor is it to be assumed that 
this relation will necessarily hold under all conditions. More- 
over, corn, notably resistant to Mg salts, does not exhibit some 
of the antagonistic effects evident in the case of wheat. No. 11 
showed pronounced chlorosis, followed, but to a somewhat less 
extent, by Nos. 7, 1, 12, 10, 13, and 14. Nos. 2-6, 8, and 15 
were normal in appearance, while No. 9 was intensely green. 

Difference in "concentration" in the first series was wholly 
in respect to a variation in the quantity of the relatively in- 
soluble salts; but inasmuch as a considerable amount of the 
insoluble residue remained in each culture at the close of the 
experiment it would seem improbable that any difference in 
the amount of the solid phase would affect the yields. Accord- 
ingly, in the series carried out at Carmel, table n and fig. 3, it 
will be seen that the following are practically the only ways in 
which the "concentrations" are varied: (1) in column "n" 
the quantity of KN0 3 is one-half the amount used in column "i," 
and (2) in column "m." while the amount of nitrate remains 



1920] 



DUGGAR — INSOLUBLE SALTS IN BALANCED SOLUTIONS 



317 




Fig. 3. Yield of wheat in solutions of relatively insoluble salts. Continuous 
line is concentration i and broken line concentration n (see table n). 



as in "i," the quantity of the iron salt is reduced to one-half. 
The concentration of KNOj was the same as in the experiments 
given in table i. The stock solution of MgS0 4 contained 41.932 
grams of the salt in 1 liter of water, so thai 10 cc. per culture 
of 240 cc. gave a solution two- thirds as strong in MgS0 4 as the 
R 6 Ci of Livingston and Tottingham ('18). In almost every 
instance where the growth quantities are sm£,ll the lower concen- 
trations of the potassium nitrate and of the magnesium sulphate 
have a tendency within the same culture number to promote the 
greater growth. In this series the best culture containing in- 
soluble salts is No. 6 (CaSO< [solid phase present!, MgNH 4 PO 
soluble ferric phosphate, and KN0 3 ), followed closely by cul- 
ture 2, the latter being the same that proved so satisfactory in 
the previous series. 

As will be pointed out later, the more insoluble calcium salts 
give the higher yields when in combination with relatively in- 
soluble salts of magnesium. This is true except in certain cases 
where iron citrate enters into the combination. In certain cul- 



*> 



318 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



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1920] 



DUGGAR — INSOLUBLE SALTS IN BALANCED SOLUTIONS 



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1920] 

DUGGAR — INSOLUBLE SALTS IN BALANCED SOLUTIONS 321 



tures where the magnesium salt is the more soluble, Nos. 8-13, 
the more favorable action of ferric citrate as contrasted with 
the soluble ferric phosphate and FeP0 4 in these cultures with 
wheat is clearly shown. With the exceptions noted the favor- 
able influence of soluble ferric phosphate in the solution is evi- 
dent, especially in Nos. 2 and 6, as also, of course, in solution B. 
The Tottingham solution was exceeded by 5 combinations. 
The Shive solution, solution A (No. 15) was unsatisfactory in 
this series, since after being set up it was found that the acidity 
was much higher than usual. In these experiments, however, 
no recrystallization of the salts employed was carried out and 

no corrections for acidity were made. 

The six cultures giving the higher yields (Nos. 2, 6, 9, 13, 16, 
and 17) were all green and healthy in appearance. Cultures 
3 and 5, without KN0 3 , were characterized by marked attenua- 
tion; No. 14 exhibited excessive greening; and Nos. 7 and 15 
were abnormally stocky in general appearance. 

As stated previously, the experiments shown in table in and 
in fig. 4 were also obtained at Carmel. The experiments were 
set up on July 10, using wheat as a test plant and employing 
quart jars as containers. For cultures 1-20 the same stock 
solutions of KN0 3 and MgS0 4 as described for table n were 
used. The results are not in entire agreement with those given 
in table n. This may be accounted for in part by the use of the 
larger containers and also in part by differences in weather 
conditions. During the progress of the experiments here dis- 
cussed, there were several days of comparatively warm weather 
without fog, inducing high evaporation rates. It is well to note 
also that a slight mishap to culture 2, which was found upset 
one morning, may be responsible in some measure for the low 

yield of this culture. 

Renewals of the solutions in the control cultures (Nos. 21- 
23) were made about every 10 days. Additional amounts of 
KNO„ 20 cc. in the case of all cultures in column "1" and 10 cc. 
in the case of column "11," were added on July 24 and August 6. 
No additional MgS0 4 was added to the cultures receiving this 
salt until August 6, when 10 cc. were given each of those receiv- 
ing this salt in column "1," and 5 cc. for similar cultures in col- 
umn "11." With the larger amount of nitrate employed, cul- 



322 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



to 



CaS0 4 -Mg 3 ( 




CaS0 4 -Mg 3 (P0 4 ) 2 - Sol # Pe.Ph.-KN0 3 
CaS0 4 -MgC03-PeP0 4 -O03 



CaS0 4 -MgC0 3 -3ol.Fe.Ph # -BI0 3 



Ca3Q4-lisNH 4 P0 4 -FeP0 4 



CaS0 4 -MgSE 4 P0 4 -PoP0 4 -003 



CaSO4-MgBH4PO.4-Sol.Pe.Ph. 



CaSO4-Mglffl4PO4-Sol.Pe.Ph.-KNO 



^ - Ca 3 (P0 4 ) 2 -MgS0 4 -5 l eP0 4 -O03 

Ca3(^04) 2 -MgS0 4 -Sol.Pe.Ph.-003 
Ca 3 (P0 4 ) 2 -MgS0 4 -Pe.Citp.-rao 3 

Ca 3 ( P0 4 ) 2 -MgS0 4 -FeC ;2 4 -EN03 

CaHP0 4 -MgS0 4 -PeP0 4 -mo 3 



CaHP0 4 -MgS0 4 -Sol.Po.Ph.-m03 



CaHPO4-MgSO4-Sol.Citr.-OO3 



CaflP04-MgS04-PeC204-QJ03 



CaC0 3 -Mg 3 1 PO4 ) E -?e80 4 -no 3 
Ca3( P0 4 ) 2 -CaS0 4 -Hg 3 ( P0 4 ) £ -3ol.F« # Hu-EHC£ 

CaHP0 4 -CaS0 4 -Mg 3 ( P0 4 ) 2 -Sol.Fe.Ph, -E3T0 3 

CaC0 3 -CaS0 4 -Mg3(P0 4 ) 2 -Sol.Fe.Ph # -003 
KH 2 P0 4 -Ca ( N0 3 ) 2 -MgS0 4 -FeP0 4 



CaS0 4 -Mg30 4 -3ol.Fe.Ph.-00 3 



KHaP0 4 -Ca( N0 3 ) s-hlgSO^-PeSOyt-rao 





Fig. 4. Yield of wheat in solutions of relatively insoluble salts. Continuous line 
is concentration 1 and broken line concentration 11 (see table in). 



19201 



DUGGAR — INSOLUBLE SALTS IN BALANCED SOLUTIONS 323 



ture 15 (CaHP0 4 , MgS0 4 , ferric citrate, and KN0 3 ) is best, 
followed by one of the controls, solution B, and this in turn is 
very closely followed by Nos. 18, 8, 20, 19, and 11. Cultures 11 
and 15 confirm the previous experience that the ferric citrate 
effects a high degree of balance in cases where the magnesium 
salt is more soluble than the calcium salt used; and neither 
the soluble ferric phosphate nor the FeP0 4 can replace it in 
this respect where wheat is the test plant (compare with the 
cultures above mentioned Nos. 9, 10, 13, and 14). 

With due consideration of the causes already mentioned the 
value of the soluble ferric phosphate in the culture media is con- 
firmed, and the importance of ferric citrate established in certain 
combinations. No experiments thus far made have thrown any 
special light on the nature of the benefit derived in these cases 
from the soluble ferric phosphate or the iron citrate. In both 
cases, however, within the range of reaction involved in the 
studies here reported, suspension films consisting in part at least 
of ferric hydroxide are thrown down. The writer is now en- 
deavoring to determine if floating particles of this type, or a 
substance in the colloidal state may possibly be of importance 
in the absorption and distribution of the ions. I have previ- 
ously mentioned this possibility (Duggar, '20, p. 42) while re- 
ferring especially to certain experiments of Bonazzi and of Allen 
on the culture of microorganisms. 



Summary 

The value of certain relatively insoluble salts as sources of 
the necessary ions for the growth of seed plants has been tested 
in a variety of combinations covering by no means, however, 
the entire range of possibility. 

It is argued that in certain types of work many advantages 
may accrue from the use of combinations of insoluble salts, 
because of (1) the tendency to maintain a constant concentra- 
tion of the various ions furnished, and also because (2) no re- 
newal of the solution (except as to the acdition of N0 3 ) is re- 
quired from day to day. 

As sources of Ca, Mg, Fe, P0 4 , S0 4 , many insoluble salts 
have been tested, but no salt of this type is procurable as a 
practical source of N0 3 , so that in most experiments this ion 
is furnished by KNO s . 



[Vol. 7 



324 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



TABLE III 



GROWTH OF WHEAT IN SOLUTIONS OF RELATIVELY INSOLUBLE SALTS. THE 
GROWTH QUANTITIES REPRESENT 8 PLANTS: PERIOD OF CULTURE, 

40 DAYS; VOLUME OF CULTURE SOLUTION, 890 cc. 



Cult. 
No. 


Concentration 




Total 

gr. wt. 
gms. 


Gr. wt. 
tops 
gms. 


Gr. wt. 
roots 
gms. 


P 


H 


i 


II 


Salts used 


Init. 


Fin. 


1 


.5 gm. 
.5 gm. 
1 . gm. 
40 cc. 




CaS0 4 +2H 2 ] 
FeP0 4 +4H 2 1 
Mg 3 (P0 4 ) 2 +8H 2 [ 
KN0 3 J 


1 11.20 


6.58 


4.62 


7.0 


7.8 


2 


.5 gm. 
1.0 gm. 
1.0 gm. 

40 cc. 




CaS0 4 +2H 2 1 
Sol. ferric phosphate 1 
Mg 3 (P0 4 ) 2 +8H 2 J 
KN0 3 J 


1 10.20 


6.01 


4.19 


8.0 


7.9 


3 


.5 gm. 
.5 gm. 
.5 gm. 
40 cc. 




CaS0 4 +2H 2 1 
FeP0 4 +4H 2 1 
MgC0 3 | 
KNO3 J 


1 9.86 


5.79 


4.07 


9.0 


7.6 


4 


.5 gm. 
1 . gm. 
.5 gm. 
40 cc. 




CaS0 4 +2H 2 1 
Sol. ferric phosphate 1 
MgC0 3 | 
KN0 3 J 


1 13.47 


8.85 


4.62 


8.8 


8.0 


5 


.5 gm.j 
.5 gm. 
.5 gm.| 


i 


CaS0 4 +2H 2 1 
FeP0 4 +4H 2 [ 
MgNH 4 P0 4 +6H20 j 


r 

1 2.40 


1.42 


.98 


7.1 


6.4 


6 


• 5 gm 
.5 gm 
.5 gm. 
40 cc. 




CaS0 4 +2H 2 1 
FeP0 4 +4Hi>0 1 
MgNH 4 P0 4 +6H 2 | 
KN0 3 J 


1 12.90 


7.72 


5.18 


6.9 


7.4 


7 


.5 gm. 

1.0 gm. 

.5 gm. 




CaS0 4 +2H 2 | 
Sol. ferric phosphate [ 
MgNH 4 P0 4 +6H 2 J 


1 5.85 


2.62 


3.23 


7 3 

1 


7.3 


8 


.5 gm. 
1.0 gm. 
.5 gm. 
40 cc. 


. 


CaS0 4 +2H 2 1 
Sol. ferric phosphate 1 
MgNH 4 P0 4 +6H 2 I 
KX0 3 J 


1 20.72 


12.05 


8.67 


8.1 


7.2 


9 


1.0 gm. 
.5 gm. 
40 cc. 
40 cc. 


1.0 gm. 
.5 gm. 
20 cc. 
20 cc. 


Ca 3 (P0 4 ) 2 ] 
FeP0 4 +4H.O 1 

MgS0 4 +7H 2 | 
KNO, J 


1 2.08 
11 8.83 


1.28 

5.77 


.80 
3.06 




7.6 

7.4 


10 


1.0 gm. 
1.0 gm. 

40 cc. 

40 cc. 


1.0 gm. 
1.0 gm. 

20 cc. 

20 cc. 


Ca 3 (P0 4 ) 2 1 
Sol. ferric phosphate 1 
MgS0 4 +7H 2 | 

KN0 3 J 


1 1.64 
11 2.32 


1.11 
1.72 


.53 
.60 


| 

8.0 


7.7 
7.7 



1920] 



DUGGAR — INSOLUBLE SALTS IN BALANCED SOLUTIONS 



325 



TABLE III— Continued 





Concentration 










Ph 


Cult. 
No. 








Total 


Gr. wt. 


Gr. wt. 








Salts used 


gr. w;. 


tops 


roots 








I 


II 




gms. 


gms. 


gms. 


Init. 


Fin. 




f 1.0 gm. 


1.0 gm. 


Ca 3 (P0 4 ) 2 1 














1.0 gm. 


1.0 gm. 


Ferric citrate 1 


1 18.02 


11.97 


6.05 


8.3 


7.9 


11 


40 cc. 
40 cc. 


20 cc. 
20 cc. 


MgS0 4 +7H 2 f 
KN0 3 J 


11 17.31 


9.86 


7.45 




8.0 




1.0 gm. 


1.0 gm. 


Ca 3 (P0 4 ) 2 1 














.5 gm. 


.5 gm. 


FeC 2 4 +2H 2 1 


1 3.48 


2.12 


1.36 


6.2 


7.5 


12 


40 cc. 
40 cc. 


20 cc. 
20 cc. 


MgS0 4 +7H 2 I 
KNO, J 


11 7. 12 


3.67 


3.45 




7.6 




.5 gm. 


.5 gm. 


CaHP0 4 +2H 2 1 














.5 gm. 


.5 gm. 


FeP0 4 +4H 2 1 


1 11.87 


7.72 


4.15 


6.6 


7.6 


13 


40 cc. 


20 cc. 


MgS0 4 +7H 2 f 


11 10.00 


6.83 


3.17 




7.5 




40 cc. 


20 cc. 


KNOs J 




1 


/ 






.5 gm. 


.5 gm. 


CaHP0 4 +2H 2 1 














1.0 gm. 


1.0 gm. 


Sol. ferric phosphate 


1 2.97 


2.07 


.90 


7.9 


7.9 


14 


40 cc. 


20 cc. 


MgS0 4 +7H 2 [ 


11 10. 67 


7.19 


3.48 




7.8 




40 cc. 


20 cc. 


KNO, 














.5 gm. 


.5 gm. 


CaHP0 4 +2H 2 ] 








1 




1.0 gm. 


1.0 gm. 


Ferric citrate 1 


1 35.^0 


22.63 


12.67 


8.1 


8.0 


15 


40 cc. 


20 cc. 


MgS0 4 +7H 2 | 


11 18.55 


11.13 


7.42 


7.3 


7.9 




40 cc. 


20 cc. 


KNO, J 


■ 




. 


1 




.5 gm. 


.5 gm. 


CaHP0 4 +2H 2 1 








f 




.5 gm. 


.5 gm. 


FeC 2 4 +2H 2 1 


1 8.80 


5.99 


2.81 


6.7 


7.9 


16 


40 cc. 


20 cc. 


MgS0 4 +7H 2 | 


11 13.14 


3.09 


6.00 




7.8 




40 cc. 


20 cc. JKNOa J 














1.0 gm. 
trace 




CaCOj 1 
FeS0 4 1 












17 


.5 gm. 




Mg 3 (P0 4 ) 2 +8H 2 I 


1 6. 52 


4.20 


2.32 


7.8 


7.9 




40 cc. 




KN0 3 J 














1.0 gm. 


1.0 gm. 


Ca 3 (P0 4 ) 2 ] 














.5 gm. 


.5 gm. 


CaS0 4 +2H 2 












18 


1.0 gm. 

.5 gm. 


1.0 gm. 
.5 gm. 


Sol. ferric phosphate \ 
Mg,(P0 4 ) 2 +8H 2 


1 25.55 


16.82 


8.73 


8.0 


7.9 




40 cc. 


20 cc. 


KNO j J 














.5 gm. 




CaHP0 4 +2H 2 | 














.5 gm. 




CaS0 4 +2H 2 












19 


1.0 gm. 
.5 gm. 




Sol. ferric phosphate \ 
Mg,(P0 4 ) 2 +8H 2 


I 18.99 


12.58 


6.41 


7.8 


8.0 




40 cc. 

■ 




KN0 8 ) 













326 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



[Vol. 7 



TABLE III— Continued 



Cult. 
No. 


Concentration 


Salts used 


Total 

gr. wt. 

gms. 


Gr. wt. 
tops 
gms. 


Gr. wt. 
roots 
gms. 


Ph 


I 


II 


Init. 


* 

Fin. 


20 


1.0 gm. 

.5 gm. 
1.0 gm. 

.5 gm. 

40 cc. 




CaC0 3 ) 
CaS04+2H 2 

Sol. ferric phosphate [ 

Mg 3 (P0 4 ) 2 +8H 2 

KNO s J 


i 19.15 


12.50 


6.65 


8.0 

1 


7.9 


21 






Solution A 


12.90 


7.92 


4.98 


4.1 


5.7 


22 






Solution B 


26.23 


15.16 


11.07 


6.6 


7.7 


23 






Tottingham's sol. 


15.55 


9.20 


6.35 


5.8 

■ 


6.4 



A relatively insoluble source of NH 4 (MgNH 4 P0 4 ) has been 
found unsatisfactory as a source of nitrogen with the test plants 
used. 

In each of three series of cultures in which wheat or wheat 
and corn were used, one or more of the combinations containing 
two or more insoluble salts exceeded the growth in the best 
control culture employed. The best control culture contained 
CaS0 4 , MgS0 4 , soluble ferric phosphate, and KNO s . Cultures 
exceeding the control contained in the several series the follow- 
ing combinations of salts: i, CaS0 4 (solid phase present), Mg» 



(poo 



2> 



soluble ferric phosphate, and KN0 3 ; n, CaS0 4 (solid 



phase present), MgNH 4 P0 4 , soluble ferric phosphate, and KNO,; 
in, CaHP0 4 , MgS0 4 , ferric citrate, and KN0 3 . 

In all series, with the test plants mentioned, a group of cul- 
tures approached very closely the yields of the best combina- 
tions, and in all cases in such best combinations the calcium 
salt is relatively more soluble than the magnesium salt, except 
in certain combinations into which ferric citrate enters. 

Soluble ferric phosphate has proved a valuable constituent 
in the culture medium in a variety of combinations. In certain 



1920] 



DUGGAR — INSOLUBLE SALTS IN BALANCED SOLUTIONS 



327 



cases ferric citrate has proved equally valuable. Certain fer- 
mentation processes may occur in cultures in which these com- 
pounds are employed and a further study of the influence of 
those changes is necessary. 

Except in the cultures containing K,P0 4 or MgCO, the hydro- 
gen-ion concentration of all combinations used in the three series 
here reported ranges from 5.6 to 8.0, and after the growth of 
test plants there is usually a shift in the P H toward alkalinity 
or greater alkalinity. 



Bibliography 

Benecke, W. ('09). Die von der Cronesche Nahrsa.zlosung. Zeitschr. f. Bot. 
1:235-252. 1909. 

Crone, G. v. d. ('03). Ergebnisse von Untersuchungen uber die Wirkung der 
Phosphorsaure auf die hohere Pflanze. Niederrheinischen Ges. f. Natur- u. 
Heilkunde, Sitzungsb. 1902; 167-173. 1903. 

Duggar, B. M. ('20). Hydrogen ion concentration and the composition of nutrient 
solutions in relation to the growth of seed plants. Ann. Mo. Bot. Gard. 7; 
1-49. /. 1-7. 1920. 

Livingston, B. E., and Tottingham, W. E. ('18). A new three-salt nutrient solu- 
tion for plant cultures. Am. Jour. Bot. 5; 337-346. 1918. 



GENERAL INDEX TO VOLUME VII 



New scientific names of plants and the final members of new combinations are 
printed in bold face type; synonyms and page numbers having reference to figures 
and plates, in italic; and previously published scientific names and all other matter, 
in ordinary type. 



A 



abietina (Hymenochaete), 186 

abietina (Thelephora) , 186 

abietinum (Stereum), 186, 246 

abnormis (Hymenochaete) , 186 

acerinum (Stereum), 236 

accrinum var. nivosum (Stereum), 236 

Acid, in relation to growth of bacteria, 

251, 258, 281, 287 
Actinomyces chromogenus, 252 
aculeata (Thelephora), 232 
affinis (Thelephora), 96 
affinis {Thelephora), 96 
albobadia (Thelephora) , 216 
albobadium (Stereum), 216, 248 
albo-marginata (Peniophora), 216 
albo-marginata (Thelephora) , 216 
Aleurodiscus subcruentatus, 237 
Alkali, in relation to growth of bacteria, 

251, 258, 281, 287 

Allen, E. R. On carbohydrate con- 
sumption by Azotobacter chroococ- 
cum, 75 

ambigua (Hymenochaete), 200 

ambiguum (Stereum), 190, 246 

ambiguus (Trichocarpus) , 200 

ambiguus (Xerocarpus), 200 

anastomosans (Stereum), 115 

anastomosans (Thelephora) , 115 

annosum (Stereum), 226 

Apparatus, Millikan dew point, 53; ro- 
tating, 76 

arcticum (Stereum), 155 

arenicolum (Stereum), 232 

areolatum (Stereum), 202 

aschistum (Corticiurn), 203 

Atkinsonii (Peniophora), 200 

aurantiaca (Podoscypha), 96 

aurantiaca (Thelephora), 75 

aurantiacum (Stereum), 95, 240 

Auricularia reflexa, 150 

australe (Stereum), 141, 244 

Autotoxin of bacteria, 250, 258, 265, 
28 1 , 287 

Azotobacter chroococcum, carbohydrate 
consumption by, 75 



B 



Bacillus aerogenes, 252, 255, in plain 
and dextrose bouillon, growth and 

Ann. Mo. Bot. Gard., Vol. 7, 1920 



hydrogen- ion concentration of, 275; 
coli, 250, in plain and dextrose bouil- 
lon, growth and hydrogen-ion concen- 
tration of, 255, 258, 281, 287; diph- 
theriae, 253; paratyphosis, 250; radi- 
cicola, 252; subtihs, 252; typhosus, 
251 

Bacterial inhibition by metabolic pro- 
ducts, 249 

Balanced solutions, insoluble salts in, 
307 

balsameum (Stereum), 145 

balsameum f . reflexum (Stereum), 145 

Beet decoction, titration curve of, 301 

Berkeleyi (Peniophora), 203 

Bertolonii (Stereum), 169 

bicolor (Stereum), 1 17 

bicolor (Thelephora), 117 

bizonatum (Stereum), 216 

Bresadolina, 81; pallida, 104 

Burtianum (Stereum), 93, 240 



c 



/ 



caespitosum (Stereum), 116, 24, 

caly cuius (Stereum), 236 
candidum (Stereum), 236 
caperata (Thelephora), 87 
caperatum (Stereum), 87, 240 
Carbohydrate consumption, by Azoto- 
bacter, 75 

carolinense (Stereum), 236 
Ceratostomella coerulea, 71 
Chailletii (Stereum), 200, 248 
Chailletii (Thelephora), 200 
Chambers, W. H. Studies in the physi- 
ology of the fungi. XI. Bacterial in- 
( hibition by metabolic products, 249 
cinerascens (Stereum), 203, 248 
cinerascens {Thelephora), 203 
cinerescens (Peniophora), 203 
Cladoderris infundibuliformis, 88 
coffearum (Stereum), 216 
coffeatum (Stereum), 117 
complicatum (Stereum), 169 
concolor (Stereum), 163 

conicum (Stereum), 179, 246 

Coniophora cerebella, 71 

Corn, food reserve in, 291; nutrition 

experiments with, 6, 310 
corrugata (Thelephora), 181 
Corticium aschistum, 203; ephebium, 204; 

lilacino-fuscum, 229; Nyssae, 128; 

(329) 



330 



INDEX 



[Vol. 7 



ochroleucum, 235 ; pezizoideum,, 121; 

siparium, 128; subrepandum, 229; 

subzonatum, 150 

Cotyledons, nutritive value of the food 

reserve in, 291 
craspedia (Thelephora), 112 

craspedium (Stereum), 113, 242 
crassa (Hymenochaete) , 192 
crassa (Thelephora), 192 
crassum (Stereum), 180 

C rater ella pallida, 104 
crateriformis (Hymenochaete), 89 
cristatum (Stereum), 103 
cristulatum (Stereum), 136 
Crone's solution, 3, 309 
Cryptochaete rufa, 121 
Culture media, liquid, titration 

of, 299 
euneatum (Stereum), 233 
cupulatum (Stereum), 179, 233 
cyphelloides (Stereum), 112, 242 
Czapek's solution, titration curve of, 301 



curves 



D 



decolor arts (Podoscypha), 107 

decolorans (Stereum), 107, 242 

decolor ans (Thelephora), 107 

diaphana (Thelephora), 98 

diaphanum (Stereum), 97, 240 

dissita (Peniophora) , 203 

dissitum (Stereum), 203 

Duggar, B. M. Hydrogen-ion concen- 
tration and the composition of nutri- 
ent solutions in relation to the growth 
of seed plants, 1 ; The nutritive value 
of the food reserve in cotyledons, 291; 
The use of "insoluble" salts in balanced 
solutions for seed plants, 307 

durum (Stereum), 226, 248 



E 



Earlei (Stereum), 199, 248 

elegans (Stereum), 93, 105, 242 
elegans (Thelephora), 105, 107 
Ellisii (Peniophora), 222 

vphebia (Peniophora), 204 
ephebium (Corticium), 204 

erumpens. (Stereum), 209, 248 

cxigua (Thelephora), 99 
exiguum (Stereum), 99, 240 



' 



F 



fasciatum (Stereum), 155, 246 

ferreum (Stereum), 202, 248 

Ferric citrate, value of, in culture media, 

321 
Ferric phosphate, soluble, value of, in 

culture media, 321 



fimbriata (Hymenochaete), 186 

fimbriatum (Stereum), 234 

fissum (Stereum), 111, 21$ 

flabellata (Podoscypha) , 111 

flabellatum (Stereum), 111 

Food reserve in cotyledons, nutritive 

value of, 291 
fragile (Stereum), 233 
frustidosa (Thelephora), 227 
frustulosum (Stereum), 227, 248 
fidvo-nitens (Stereum), 91 
Fungi, studies in the physiology of the, 

XI, 249 
fusca (Thelephora), 117 

fuscum (Stereum), 117, 244 



G 



Galeottii (Stereum), 234 

gausapata (Thelephora), 136 
gausapatum (Stereum), 136, 244 
Geographic distribution of Stereums, 82 
glabrescens (Stereum), 110, 242 
glaucescens (Stereum), 186 
griseum (Stereum), 234 
Growth curves of bacteria, 249 
Growth of : Azotobacter chroococcum, 75 ; 
corn in nutrient solutions, 6, 310; 
peas in nutrient solutions, 6; wheat 
in nutrient solutions, 6, 309; seedlings, 
effects of cotyledons upon, 291 
guadelupense (Stereum), 236 



H 



Hartmanni (Stereum), 112, 242 

Hartmanni (Thelephora), 112 

Haydeni (Stereum), 236 

Helvetia pannosa, 104; versicolor, 167 

heterosporum (Stereum), 220, 248 

hirsuta (Thelephora), 150 

hirsuta var. ramealis (Thelephora), 169 

hirsutum (Stereum), 150, 246 

Huberianum (Stereum), 111 

Humidity, in relation to moisture imbi- 
bition by wood and to spore germina- 
tion on wood, 51; determination of, 

53 

Humidors used in wood-decay experi- 
ments, description of, 52; view of, 74 

Hydrogen-ion concentration, and the 
composition of nutrient solutions in 
relation to the growth of seed plants, 
1, 327; in relation to growth of bac- 
teria, 252, 256, 258, 281, 287 

hydrophorum (Stereum), 89, 240 

Hymenochaete abietina, 186; abnarmis, 
186; ambigua, 200; crassa, 192; crateri- 
formis, 89; fimbriata, 186; Kalchbren- 
neri, 192; multispinulosa, 192; pallida, 
196; pauper cula, 216; purpurea, 192; 









1920] 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



331 



ruqispora, 188; scabriscta, 192; um- 
brina, 192; vinosa, 192 
Hypocrea Richardsonii, 121 



I 



illudons (Stereum), 225 

Imbibition by wood, relation of, to hu- 
midity, 51 

infundibuliforrnis (Cladodcrris), 88 
insigne (Stereum), 225, 248 
insolitum (Stereum), 237 

Insoluble" salts in balanced solutions, 
307 

intermedia (Peniophora), 192 



a 



K 

Kalchbrenneri (Hymenochaete), 192 
Karrer, J. L., and R. W. Webb. Titra- 
tion curves of certain liquid culture 
media, 299 

Kneiffia purpurea, 192 



L 



lamellate (Thelephora) , 88 
lamellatum (Stereum), 88 

Lenzites saepiaria, humidity and spore 

germination of, 51 
Leveillianum (Stereum), 237 
lilacino-juscum (Corticium), 229 
lilacino-fuscum (Stereum), 229 
Lloydella, 81; occidentalis , 204; 

riscta, 192; striata, 186 
lobata (Thelephora), 163, 169 

lobatum (Stereum), 163, 246 



scab- 



M 

macrorrhiza (Thelephora), 93 
macrorrhizum (Stereum), 92 
magnisporum (Stereum), 207, 248 

Mancianus (Stereum), 237 

Media, liquid culture, titration curves 
of, 299 

Merulius lacrymans, 71 

Metabolic products, bacterial inhibition 
by, 249 

Micheneri (Stereum), 128, 237 

Mineral nutrient solutions, 1, 307 

Moisture content of wood, 54; deter- 
mination of, 54 

molle (Stereum), 155 

mollis (Thelephora), 155 

moricola (Peniophora), 203 

moricola (Stereum), 203 

multispinulosa (Hymenochaete) , 192 

Murraii (Thelephora), 131 

Murrayi (Stereum), 131, 244 

mytilina (Thelephora), 141 



N 

neglecta (Peniophora), 204 
neglectum (Stereum), 204 
nicaraguense (Stereum), 196 
nitidulum (Steieum), 101 
Nutrient solutions for seed plants, 
Nyssae (Corticiim), 128 



1,307 



o 



obscura (Peniophora), 222 
obscura (Thelephora), 222 
occidentale (Stereum), 136 
occidentalis (Lloydella), 204 
occidentalis (Peniophora), 20-4 
ochracea (Thelephora), 150 
ochraceo-flava (Thelephora), 183 
ochraceo-flavuri (Stereum), 183, 246 
ochroleucum (C \rrticium) , 235 
ochroleucum (Stereum), 148, 235 
ostrea (Stereum;, 155 
ostrea (Thelephora), 155 



196 



24 






p 



pallida (Bresadolina) , 104 
pallida (Craterella), 104 
pallida (Hymenochaete), 
pallida (Thelepnora), 104 
pallidum (Stereum), 104, 
pannosa (Helvetia), 104 
pannosa (Thelephora), 104 
papyrina (Peniophora), 196 
papyrinum (Stereum), 196, 248 

patelliforme (Stereum), 182, 246 

paupercula (Hymenochaete), 216 

pauper cula (Peniophora), 216 

Peas, Canada f eld, food reserve of, 291; 
nutrition experiments with, 6 

Peniophora albo-marginata, 216; Atkin- 
sonii, 200; Berkeleyi, 203; cinerescens, 
203; dissita, 203; Ellisii, 222; ephebia, 
204; intermedia, 192; moricola, 203; 
neglecta, 204; obscura, 222; occiden- 
talis, 204; papyrina, 196; paupercula, 
216; Schweinltzii, 203 

Peptone solution, titration curve of, 

perdix (Thelephora), 227 

pergamenum (Stereum), 101, 240 

petalodes (Stereum), 114, 242 

pezizoidea (Tuberculoma) , 121 

pezizoideum (Corticium), 121 

Pfeffer's solution, titration curve of, 301 
Pini (Sterellum), 123 
Pini (Stereum) 123, 244 
Pini (Thelephora), 123 
Pinus echinata, imbibition 

51; palustris, imbibition 

51 

Pneumococcus, 253 



301 



of 
of 



wood 
wood 



of, 
of, 



[Vol. 7 



332 



INDEX 



Podoscypha, 81; aurantiaca, 96; decolor- 

arts, 107; flabellata, 111; radicans, 108 
populneum (Stereum), 237 
prolifera (Thelephora), 115 
proliferum (Stereum), 115, 244 
pruinatum (Stereum), 237 
pubescens (Stereum), 178, 246 

pulvendentum (Stereum), 131 
purpurascens (Stereum) , 204 
purpurea (Hymenochaete), 192 
purpurea (Kneifia), 192 
purpurea (Thelephora), 124 
purpureum (Stereum), 124, #44 
pusiolum (Stereum), 109, 242 




quisquiliare (Stereum), 95, 240 
quisquiliaris (Thelephora), 95 



R 



radians (Stereum), 167 
radiatum (Stereum), 181, 246 
radiatum var. reflexum (Stereum), 181 
radicans (Podoscypha), 108 
radicans (Stereum), 108, 242 
radicans (Thelephora), 108 
rameale (Stereum), 169, 246 
Ravenelii (Stereum), 90, 240 
reflexa (Auricularia) , 150 
Resin content of wood, determination of, 
54; in relation to moisture absorption, 

66 
Richards' solution, titration curve of, 

301 

Richardsonii (Hypocrea), 121 

rigens (Stereum), 145 
rivulorum (Stereum), 94 
roseo-carnea (Thelephora), 229 
roseo-carneum (Stereum), 229, 248 
rufa (Cryptochaete), 121 
rufa (Thelephora), 121 

rufum (Stereum), 120, 244 
rugispora (Hymenochaete), 188 
rugisporum (Stereum), 188, 248 
rugosa (7 1 helcphora) , 143 

rugosiusculum (Stereum), 127, 244 
rugosum (Stereum), 142, 244 



s 



Salt requirements of seed plants, 1 
sanguinolenta (Thelephora), 145 
sanguinolentum (Stereum), 144, 246 
saxitas (Stereum), 134, 244 
scabriseta (Hymenochaete), 192 
scabriseta (Lloydella), 192 
Schweinitzii (Peniophora), 203 
scriblitum (Stereum), 237 
Seed plants, hydrogen ion concentration 
and the composition of nutrient solu- 



tions in, 1; "insoluble" salts in bal- 
anced solutions for, 307 
sendaiense (Stereum), 229 

sepium (Stereum), 215, 248 
seriatum (Stereum), 237 
sericella (Thelephora), 96 
sericeum (Stereum), 175, 246 
Shive's solution, 3 
siparium (Corticium), 128 
Solutions, balanced, "insoluble" Baits 
in, for seed plants, 307 

Sowerbeyi (Stereum), 104 
Sowerbeyi (Thelephora), 104 
spadicea (Thelephora), 136 
spadiceum (Stereum), 136 
spadieeum var. plicatum (Stereum), 136 

spectabilis (Thelephora), 96 
spongiosum (Stereum), 237 
Spore germination on wood, 68 
Sprucei (Stereum), 163 

spumeum (Stereum), 208, 236, 248 

Sterellum Pini, 123 

Stereum, 81; abietinum, 186, 246; accr- 
inum, 236, var. nivosum, 236; albo- 
badium, 216, 248; ambiguum, 190, 
246; anastomosans, 115; annosum, 
226; arctiewn, 155; arenicolum, 232; 
areolatum, 202; aurantiacum, 95, 240; 
australe, 141, 244; balsameum, 145, 
f. reflexum, 145; Bertolonii, 169; bi- 
color, 117; bizonatum, 216; Burtianum, 

93, 240 ; caespitosum, 116, 244;caly- 

culus, 236; candidum, 236; caperatum, 
87, 240 ; carolinense, 236; Chailletii, 
200, 248; cinerascens, 203, 248; cof- 
fcarum, 216; coffeatwn, 117; cow pli- 
catum, 169; concolor, 163; conic um, 
179, 246; craspedium, 113, 242; eras- 
sum, 180; cristatum, 103; cristulatum, 

136; cuneatum, 233; cupulatum, 179, 
233; cyphelloides, 112, 242; decolor- 
ans, 107, 242; diaphanum, 97, 240; 
dissitum, 203; durum, 22$, 248; Earl- 
ei, 199, 248; elegans, 93, 105, £&; 
erumpens, 209, 248;exig\ium, 99,240; 
fasciatum, 155, 246 ;icrreum, 202, 248; 
fimbriatum, 234; fissum, 111, 242; 
flabellatum, 111; fragile, 233; frustu- 
losum, 227, 248; fulvo-nitens, 91; fus- 
cum, 117, 244; Galeottii, 234; gausa- 
patum, 136, 244; glabrescens, 110, 
242; glaucescens, 186; griscum, 234; 
guadelupense, 236; Hartmanni, 112, 
242; Haydeni, 236; heterosporum, 

220, 248; hirsutum, 150, 246; Hubert- 
anum, 111; hydrophorum, 89, 240; 
illudens, 225; insigne, 225, 248; in- 
solitum, 237; lamellatum, 88; Leveilli- 
anum, 237; lilacino-fuscum, 229; lo- 
batunij 163, 246; macrorrhizum, 92; 
magnisporum, 207, 248; Mancianm, 

237; Micheneri, 128, 237; molle, 155; 
moricola, 203; Murrayi, 131, 244 > wg- 



1920] 



ANNALS OF THE MISSOURI BOTANICAL GARDEN 



333 



ledum, 204; nicaragven.se, 196; nitidu- 
lum, 101; occidentale , 136; ochraceo- 
flavum, 183, 246; ochroleucum, 148, 
235; ostrea, 155; pallidum, 104, 242; 

papyrinum, 196, 248; patelliforme, 
182, 246; pergamenum, 101, 240; peta- 
lodes, 114, 242; Pini, 123, 244; popul- 
neum, 237; proliferum, 115, 244 
pruinatum, 237 ; pubescens, 178, 246 
pulverulentum, 131; purpurascens, 204 
purpureum, 124, 244; pusiolum, 109 
242; quisquiliare, 95, 240; radians, 167 
radiatum, 181, 246; radiatum var. re- 
flexum, 181; radicans, 108, 242; rame- 
ale, 169, 246; % Ravenelii, 90, 240; 
rigens, 145; rivulorum, 94; roseo- 
carneum, 229, 248; rufum, 120, 244; 
rugisporum, 188, 248; rugosiusculum, 
127, 244; rugosum, 142, 244; sanguino- 
lentum, 144, 246; saxitas, 134, 244; 
scriblitum, 237; sendaiense, 229; se- 
pium, 215, 248; seriatum, 237; seri- 
ceum, 175, 246; Sowerbeyi, 104; spadi- 
ceum, 136, var. plicatum, 136; spongi- 
osum, 237; Sprucei, 163; spumeum, 
208, 236, 248; strumosum, 237; sty- 
racifluum, 135, 244; subcruentatum, 
237; subpileatum, 213, 248; sulcatum, 
211, 248; Sidlivantii, 98; sulphuratum, 
148, 246; surinamense, 91, 240; tener- 
rimum, 100, 240; triste, 238; tuber- 
culosum, 131; umbrinum, 191, 248; 
undulatum, 100; unicum, 236; varii- 
color, 150; versicolor, 166, 246; versi- 
forme, 222, 248; vibrans, 179, 246; 
vorticosum, 124; Willeyi, 98; xanthel- 

lum, 96 

Streptococcus erysipelatus, 252; hemo- 
lytics, 253; lacticus, 252 

striata (Lloydella), 186 

striata (Thelephora), 186 

striatum (Stereum), 186 

strumosum {Stereum), 237 

Studies in the physiology of the fungi. 

XI, 249 

styraczflua (Thelephora), 135 
styracifluum (Stereum), 135, 244 
subcruentatum (Stereum), 237 
subcruentatus (Aleurodiscus), 237 
subpileatum (Stereum), 213, 248 
subrepandum (Corticium), 229 
subzonata (Thelephora), 150 
subzonatum (Corticium), 150 
sulcatum (Stereum), 211, 248 
Sullivantii (Stereum), 98 
sulphuratum (Stereum), 148, 246 

surinamense (Stereum), 91, 240 

T 

tenerrimum (Stereum), 100, 240 
Thelephora abietina, 186; aculeata, 232; 
affinis, 96; afinis, 96; albobadia, 216; 
a tbo-marginata, 216 ; anastomosans , 
115; aurantiaca. 75; bicolor, 117; caper- 
ata, 87; Cliallletii, 200; cinerascens, 



203; corrugata, 181; craspedia, 112; 
crassa, 192; decolorans, 107; diaphana, 
98; elegans, 105, 107; exigua, 99; 
frustidosa, 227;fusca, 117; qausapata, 
136; Hartmanni, 112; hirsuta, 150, var. 
ramealis, 16S; lamellata, 88; lobata, 
163, 169; macrorrhiza, 93; mollis, 155; 
Murraii, 131; mytilina, 141; obscura, 
222; ochracea, 150; ochracea-flava, 183; 
0£traz, 155; pallida, 104; pannosa, 104; 
pento, 227; Ptfii, 123; prolifera, 115; 

purpurea, 124; quisquiliaris, 95; 
radicans, 108; roseo-carnea, 229; ri//a, 
121; rugosa, 143; sanguinolenta, 145; 
sericella, 96; Sowerbeyi, 104; spadicea, 
136; spectabUis, 96; s<nato, 186; $<?/- 
raciflua, 135; subzonata, 150; trem- 
coiar, 167, var. fasciata, 155 

Titration curves, culture media, 299 

Trichocarpus ambiguus, 200 

triste (Stereum] , 238 
Tuber cularia peiizoidea, 121 
tuberculosum (Stereum), 131 

u 

umbrina (H yme nochaete) , 192 

umbrinum (Stereum), 191, &4S 
undulatum (Stereum), 100 

unicum (Stereu :n), 236 

v 

variicolor (Stereum), 150 
versicolor (Helvetia), 167 
versicolor (Stereum), 166, &}# 
versicolor (Thelephora), 167 
versicolor var. fjsciata (Thelephora), 
versiforme (Stereum), 222, j&$8 
vibrans (Stereum), 179, &$0 
vinosa (Hymencchaete), 192 
Vitamine, in relation to cotyledons, 
vorticosum (Stereum), 124 

w 

Webb, R. W., J. L. Karrer, and. Titra- 
tion curves n certain liquid culture 

media, 299 
Wheat, nutrition experiments with, 6, 

309 

Willeyi (Stereum), 98 

Wood, imbibition in relation to humid- 
ity of, 51 

Wood-destroying fungi, moisture in re- 
lation to, 51 

x 

xanthellum (St3reum), 96 
Xerocarpus aminguus, 200 



155 



297 



z 

Zeiler, S. M. Humidity in relation 
moisture imbibition by wood and 
spore germination on wood, 51 



to 
to